A polytypic species revisited: Phylogenetic and morphological variation, taxonomic status, and geographical distribution of Trachops (Chiroptera: Phyllostomidae)

<table><tr><td>Journal:</td><td>Journal of Mammalogy</td></tr><tr><td>Manuscript ID</td><td>Draft</td></tr><tr><td>Manuscript Type:</td><td>Research Article</td></tr><tr><td>Date Submitted by the Author:</td><td>n/a</td></tr><tr><td>Complete List of Authors:</td><td>Camacho, M. Alejandra; Pontificia Universidad Católica del Ecuador, Museo de Zoología Menéndez-Guerrero, Pablo; Pontificia Universidad Católica del Ecuador, 3Laboratorio de Macroecología y Cambio Global, Facultad de Ciencias Exactas y Naturales Horváth, Balázs; Bernhard Nocht Institute of Tropical Medicine, WHO Collaborating Centre for Arbovirus and Haemorrhagic Fever Reference and Research Cadar, Dániel; Bernhard Nocht Institute of Tropical Medicine, WHO Collaborating Centre for Arbovirus and Haemorrhagic Fever Reference and Research Murienne, Jérôme; Université Toulouse III-Paul Sabatier, Centre de Recherche sur la Biodiversité et l'Environnement (CRBE)</td></tr><tr><td>Keywords:</td><td>Fringed-Lipped Bat, Mitochondrial DNA analysis, Neotropical Phyllostomids, Taxonomic evaluation, T. cirrhosus, T. ehrhardti, Trachops diversity</td></tr></table>
A polytypic species revisited: Phylogenetic and morphological variation, taxonomic status, and geographical distribution of *Trachops* (Chiroptera: Phyllostomidae)

* M. Alejandra Camacho¹,² 0000-0001-7965-184X, Pablo A. Menéndez-Guerrero³ 0000-0002-3276-4537, Balázs Horváth⁴ 0009-0000-4632-1908, Dániel Cadar⁴ 0000-0002-8063-2766, Jérôme Murienne¹ 0000-0003-1474-7829

¹Centre de Recherche sur la Biodiversité et l'Environnement (CRBE), Université de Toulouse, CNRS, IRD, Toulouse INP, Université Toulouse 3 - Paul Sabatier (UT3), Toulouse, France.

²Museo de Zoología, Facultad de Ciencias Exactas y Naturales, Pontificia Universidad Católica del Ecuador, Quito, Ecuador.

³Laboratorio de Macroecología y Cambio Global, Facultad de Ciencias Exactas y Naturales, Pontificia Universidad Católica del Ecuador, Quito, Ecuador

⁴WHO Collaborating Centre for Arbovirus and Haemorrhagic Fever Reference and Research, Bernhard Nocht Institute for Tropical Medicine, 20359 Hamburg, Germany

*Corresponding author: M. Alejandra Camacho. macamachom@puce.edu.ec, Av. 12 de Octubre 1076 y Roca. Zip Code: 170525.

**Short running title:** Phylogenetic and morphological characterization of *Trachops*

**Abstract**

The taxonomic status of the neotropical bat genus *Trachops* has been reevaluated through an integrated study that incorporates morphological, morphometric, and molecular data across its extensive geographic range. Our research, which included previously unexamined regions, revealed substantial insights into the diversity within *Trachops*. The results support the elevation
of *T. cirrhosus ehrhardti* to species status, due to genetic and morphological differences in southeastern Brazil specimens. Conversely, our comprehensive analysis found insufficient evidence to maintain the subspecific distinction of *T. c. coffini*, which lacks diagnosable morphological characters and is not genetically distinct from *T. c. cirrhosus* across its distribution range. Additionally, our findings challenge the previous belief of a latitudinal differentiation in body size for *Trachops cirrhosus*, as specimens from western South America and northeastern South America exhibit similar sizes to those from Central America. These results underscore the importance of revising the taxonomic framework for this bat genus, contributing to a more precise understanding of its evolutionary relationships, and further enhancing conservation efforts, considering the potential threats to the newly recognized *T. ehrhardti* in the imperiled Atlantic Forest of Brazil.

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**Keywords:** Fringed-Lipped Bat, Mitochondrial DNA analysis, Neotropical Phyllostomids, Taxonomic evaluation, *T. cirrhosus*, *T. ehrhardti*, *Trachops* diversity.

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## 43 Resumen

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El estatus taxonómico de los murciélagos neotropicales del género *Trachops* ha sido reevaluado mediante un estudio integrado que incorpora datos morfológicos, morfométrico y moleculares a través de su extensa distribución geográfica. Nuestra investigación, que incluyó regiones previamente no examinadas, reveló perspectivas sustanciales sobre la diversidad dentro del género *Trachops*. Los resultados respaldan la elevación de *T. cirrhosus ehrhardti* al estatus de especie debido a las diferencias genéticas y morfológicas en especímenes en el sudeste de Brasil. Por otro lado, nuestro análisis no encontró suficiente evidencia para mantener la distinción subespecífica de *T. c. coffini*, que carece de caracteres morfológicos diagnosticables, y no difiere genéticamente de *T. c. cirrhosus* en su rango de distribución. Adicionalmente, nuestros hallazgos confrontan las creencias previas de una diferenciación latitudinal en el tamaño corporal de
55 *Trachops cirrhosus*, ya que especímenes del oeste y noreste de Sudamérica exhiben tamaños similares a los encontrados en individuos de Centroamérica. Estos resultados subrayan la importancia de examinar el marco taxonómico para este género de murciélagos, contribuyendo a una comprensión más precisa de sus relaciones evolutivas, para aumentar los esfuerzos de conservación a futuro, considerando los riesgos potenciales a la recientemente reconocida especie
60 *T. ehrhardti* en los Bosques Atlánticos amenazados de Brasil.

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62 **Palabras clave:** Análisis DNA mitocondrial, diversidad *Trachops*, Evaluación taxonómica, Filostómidos Neotropicales, Murciélago de labio verrugoso, *T. cirrhosus*, *T. ehrhardti*

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65 **Introduction**
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The Frog-Eating bat, *Trachops cirrhosus* (Spix 1823), is a member of the family Phyllostomidae (Gray 1825), and the only member of its genus (Solari et al. 2019). This species has a wide distribution in the Americas, ranging from southern Mexico to Brazil, and occurring at mid to high elevations at both sides of the Andes Cordillera, across the Amazon, and in the Atlantic Forest (Jones and Carter 1976; Cramer et al. 2001; Williams and Genoways 2008; Solari et al. 2019). The species occurs in humid tropical and subtropical forests, and its distribution encompasses primary, secondary, disturbed and gallery forests, at woodland edge and near cultivated areas (Fenton et al. 1992; Ditchfield 1996; Tirira 2017; Solari et al. 2019).

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*Trachops* is monophyletic genus (Ditchfield 1996; Baker et al. 2003; Camacho et al. 2022 and can be easily distinguished from other bats by its finger-like dermal projections on the chin and lips (Figure 1). Historically, *Trachops* taxonomy has undergone numerous changes (Spix 1823; Gray 1825, Gray, 1847; Felten 1956a; Schinz 2012) until the late 1950s, when its current taxonomy was stablished with three recognized subspecies: *Trachops cirrhosus cirrhosus* (Spix 1823), occurring from Costa Rica to northeastern Brazil; *Trachops cirrhosus coffini* (Goldman 1925), distributed
from Mexico to Nicaragua; and *Trachops cirrhosus ehrhardti* (Felten, 1956b), which is found only in southeastern Brazil (Figure 2) ([Jones and Carter 1976; Solari et al. 2019]).

The descriptions of the subspecies of *Trachops cirrhosus* are brief and lack comparisons with specimens from Central and South America locations, leading to unclear geographic ranges and diagnostic characters. Moreover, only a few recent studies have addressed the systematics of *Trachops* itself. Ditchfield (1996) used partial sequences of the mitochondrial Cytochrome b gene and identified 5 distinct haplotypes with an average sequence divergence ranging from 5.5% to 11%, demonstrating strong geographic structuring and limited sharing of haplotypes among distant localities. Later, and despite the high genetic divergences yielded by the mitochondrial marker COI, Clare et al. (2007) and Clare (2011) could not find congruence with the topology yielded by the nuclear marker Dby. However, they observed many intraspecific groups occupying sympatric distributions, strongly suggesting ongoing or past speciation events. Most recently, Fonseca (2019) proposed elevating *T. c. ehrhardti* to the species level, based on integrative evidence in morphology, ecology, and genetics, while maintaining *T. c. cirrhosus* and *T. c. coffini* as subspecies of *T. cirrhosus*. But these taxonomic changes, initially proposed by Fonseca (2019), have not yet been formalized in a peer-reviewed publication, and the morphological guidelines for characterization of *Trachops cirrhosus* subspecies remain unclear.

In the present study, we revise the taxonomic composition and phylogenetic structure of the *Trachops* genus based on molecular, morphological, morphometric, and geographic data that includes a significant sample from the West South American region. We aim to address this issue by answering the following questions: (1) Do the current subspecies of *Trachops cirrhosus* have valid phylogenetic and morphological justifications? (2) What are the distinct and diagnosable characters of each subspecies? (3) Can biodiversity of cryptic species be inferred in *Trachops* through molecular and morphological analysis? We seek to clarify, correctly diagnose, and geographically limit *T. cirrhosus* subspecies. We used a phylogenomic dataset based on complete
mitochondrial genome which has provided meaningful information in Phylostomids (Camacho et al. 2022). In addition, we performed an extended taxon sampling, ensuring comprehensive spatial coverage. This diverse geographic sampling encompasses a wide range of habitats and ecological zones where *Trachops cirrhosus* subspecies are found.

## Methods

**Specimens, tissue samples, and biorepositories:** For the morphometric analyses, we measured 238 specimens of *Trachops cirrhosus*. Only adults were included, following the consideration from Kunz et al. (1996) for sex, age, and reproductive conditions of mammals. We included representatives from all named subspecies and from their complete distribution areas in Central and South America. The samples comprised fluid-preserved specimens, study skins, and skulls deposited in the following institutions: **AMNH**, American Museum of Natural History, New York, NY, USA; **IAvH**, Colección de Mamíferos del Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Villa de Leyva, Colombia; **MEPN**, Museo de Historia Natural Gustavo Orcés, Escuela Politécnica Nacional, Quito, Ecuador; **QCAZ**, Museo de Zoología, División de Mastozoología, Pontificia Universidad Católica del Ecuador, Quito, Ecuador; **SMF**, Senckenberg Naturmuseum Frankfurt, Frankfurt, Germany; **USNM**, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; and **UV**, Colección del Mamíferos de la Universidad del Valle, Cali, Colombia. The measurements of the specimens of *Trachops cirrhosus* ehrhardti from the SMF (n = 3) were shared by the curator of this collection. We obtained tissue samples from the following collections: **AMNH**; **FMNH**, Field Museum of Natural History, Chicago, IL, USA; **MSB**, Museum of Southwestern Biology, Albuquerque, NM, USA; **QCAZ**; **ROM**, Royal Ontario Museum, Toronto, Canada; and **SMF** (Supplementary Table 1).

**Genomic DNA isolation, amplification, and sequencing:** We gathered a dataset comprising 78 tissue samples, some of them dating back to 1910. The tissue samples encompassed heart, liver,
claws, and wing snippets. Mitochondrial DNA sequencing was achieved using a genome skimming procedure following Camacho et al. (2022). Laboratory procedures were carried out at the NGS Core Facility of the Bernhard Nocht Institute for Tropical Medicine in Hamburg. The extraction and amplification of archival DNA were performed in a dedicated clean room facility, which was separate from the area where current samples and post-PCR products were handled. Stringent contamination prevention protocols and negative controls were also implemented.

The DNA extraction process for various sample types (dried skin, ethanol, or formaldehyde-preserved tissues) involved proteinase K digestion at 55°C using 20 µl of proteinase K and 220 µl of ATL lysis buffer (MinElute Reaction Cleanup kit, Qiagen). Prior digestion, the samples were thoroughly washed with nuclease-free water (Qiagen). The incubation time for proteinase K digestion varied depending on the tissue type and sample preservation, ranging from 5 to 24 hours. Following digestion, DNA was extracted and purified using the Qiagen MinElute kit, with each sample eluted to a final volume of 60 µl. DNA concentration was measured using Qubit and Bioanalyzer instruments. For library preparation, the QIAseq FX DNA Library Kit (Qiagen) was used, with double index barcode labelling according to the manufacturer's instructions. DNA fragmentation was often avoided due to the high degradation of nucleic acid material and low concentration. The HiFi PCR Master Mix from the QIAseq FX kit was utilized to amplify DNA regions with varying GC contents, minimizing sequencing bias caused by PCR, such as nucleotide misincorporations from cytosine deamination. Subsequently, the libraries underwent quality control to determine fragment size using the Agilent 2100 Bioanalyzer, and concentration was assessed using a Qubit 2.0 Fluorometer. After normalization, the samples and negative controls were pooled and subjected to sequencing on the NextSeq 2000 platform (2 × 100 cycles) (Illumina, San Diego, CA, USA).

**Mitogenomes assembly and annotations:** Raw reads were first subjected to a qualitative assessment, followed by the removal of adaptor sequences and the filtration of polyclonal and
163 low-quality reads (<55 bases long) using CLC Workbench (Qiagen). Overlapping paired-end (PE) reads were merged to improve quality, while non-overlapping pairs and orphan reads were left unchanged. Deduplication was performed with an assumed 100% identity using BBTools (Bushnell 2014), expanding the length of contigs produced during de novo assembly. Custom assembly was conducted using Megahit (Li et al. 2016) and Spades (Prjibelski et al. 2020) applications. A specialized Chiroptera mitochondrial database and BLASTN were employed to identify potential bat mitochondrial genomes in the resulting contigs. This process was followed by remapping and visual validation of circularization using CLC Genomics Workbench 22. Finally, the GeSeq online tool (Tillich et al. 2017) was used to annotate genomic features. All the assembled mitochondrial genomes were annotated using the MITOS2 metazoan pipeline (Bernt et al. 2013; Al Arab et al. 2017), followed by manual adjustment in Geneious v.9.0.5 (https://www.geneious.com). The mitochondrial DNA sequences obtained in this study have been deposited in GenBank (Appendix 1).

176 **Phylogenetic analysis:** Ribosomal RNA (rRNA) and transfer RNA (tRNA) loci were aligned using MUSCLE (Edgar 2004), while protein-coding genes sequences were aligned using TranslatorX (Abascal et al. 2010). As outgroups, we selected sequences from *Phyllostomus hastatus*, *Macrophyllum macrophyllum*, and *Phylloderma stenops*, shown as closely related in previous phylogenetic studies (Botero-Castro et al. 2018; Camacho et al. 2022). We used the optimal partitioning scheme of 38 partitions (2 independent partitions for rRNAs and 36 partitions for tRNAs, one partition for each codon for each gene) and a Generalized Time Reversible (GTR) model of substitution rates along with a gamma (G) distribution and a fraction of invariable (I) sites (GTR+G+I).

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186 We performed a Maximum Likelihood (ML) analysis using RAxML-NG (Kozlov et al. 2019), starting from 10 parsimony trees and 10 random trees. Bootstrap support values were obtained using the classical Felsenstein metric (Felsenstein 1985) and transfer bootstrap expectation (Lemoine et al. 2018). Bayesian inference (BI) analyses were performed using MrBayes v.3.2.7
190 (Ronquist et al. 2012). We partitioned the sequences in 38 sets corresponding to 2 independent partitions for rRNAs and for tRNAs plus 36 partitions, one partition for each codon for each gene, and used the best analytical scheme as evaluated by the AIC. We ran 8 Markov chain Monte Carlo (MCMC) chains for 10 million generations, with default heating values. The sampling frequency was set every 1000 generations, and the first 25 000 samples were discarded as burn-in. A consensus tree was built under the majority rule consensus of all trees obtained in the 8 runs after the burn-in period. We used the 'sumt' command to produce summary statistics for trees sampled during a Bayesian MCMC analysis. Posterior probabilities of nodes were regarded as estimators of confidence. Trees were visualized and edited in FigTree v.1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/). We adopted the nodal support criteria established by Moratelli et al. (2017): in the ML analysis, robust support is considered when bootstrap values exceed or equal to 75%, while support is deemed moderate when values range from 50% to 75%. Values equal to or below 50% are indicative of negligible support in our analyses. We calculated uncorrected pairwise (p) distances within and among samples of *Trachops cirrhosus* using MEGA 11 (Stecher et al. 2020; Tamura et al. 2021).

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206 **Morphological and morphometric analyses:** A total of 27 measurements were taken. Among these, 9 cranial measurements and 4 external measurements were selected for statistical analyses, as indicated by asterisks (*) below; the remaining measurements were employed for descriptive purposes:

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211 *Calcar length (CL)*: From the joint with the ankle to the calcar tip.

212 *Ear length (E)*: Intertragic notch of the ear to the outer tip

213 *Forearm length (FA)*: Distance from the elbow (tip of the olecranon process) to the wrist (including the carpals). This measurement is taken with partially folded wings.

215 *Hindfoot length (HF)*: Distance from the ankle to the tip of the claw.
216 *Metacarpal III (MET-III): Distance from the joint of the wrist (carpal bones) with the 3rd metacarpal to the metacarpophalangeal joint of 3rd digit.

218 *Metacarpal IV (MET-IV): Distance from the joint of the wrist (carpal bones) with the 4th metacarpal to the metacarpophalangeal joint of 4th digit.

220 *Metacarpal V (MET-V): Distance from the joint of the wrist (carpal bones) with the 5th metacarpal to the metacarpophalangeal joint of 5th digit.

222 Tail length (T): Distance from dorsal flexure at base of the tail to the tip of the last caudal vertebra.

223 Tibia length (TiL): Length from the proximal end of the tibia to the distal base of the calcar.

224 Total length (TL): Head and body length excluding tail.

225 Weight (W): Mass in grams.

226 Braincase height (BCH): Height of the braincase, posterior to the auditory bullae from the basioccipital to the sagittal crest.

228 *Breadth across upper molars (M2-M2): Greatest width of palate across labial margins of the alveoli of M2s.

230 *Breadth of brain case (BB): Greatest breadth of the globular part of the braincase, excluding mastoid and paraoccipital processes.

232 Condylocanine length (CCL): Distance from the occipital condyles to the anterior border of the upper canines.

234 *Condyloincisive length (CIL): Distance between a line connecting the posteriormost margins of the occipital condyles and the anteriormost point on the upper incisors.

236 Coronoid height (COH): Perpendicular height from the ventral margin of mandible to the tip of coronoid process.

238 Dentary length (DENL): Distance from midpoint of condyle to the anteriormost point of the dentary.

240 *Greatest length of skull (GLS): Greatest distance from the occiput to the anteriormost point on the premaxilla (including the incisors).
242 *Mandibular tooth throw length (MANDL)*: Distance from the anteriormost surface of the lower canine to the posteriormost surface of m3.

244 *Mastoid (process) breadth (MPW)*: Greatest breadth across skull, including mastoid processes.

245 *Maxillary tooth throw (MTRL)*: Distance from the anteriormost surface of the upper canine to the posteriormost surface of the crown of M3.

247 *Molariform tooth throw (MLTRL)*: Distance from the anteriormost surface of P3 to the posteriormost surface of the crown of M3.

249 *Palatal width at canines (C-C)*: Distance between the outermost extremities of the cinguli of upper canines.

251 *Palatal length (PL)*: Distance from the posterior palatal notch to the anteriormost border of the incisive alveoli.

253 *Post orbital constriction breadth (PB)*: Least breadth at the postorbital constriction.

254 *Zygomatic breadth (ZB)*: Greatest breadth across the zygomatic arches.

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256 External and osteological characteristics were based on, but not limited to the guidelines proposed by Velazco (2005), Tavares et al. (2014), Molinari et al. (2017), and Garbino et al. (2020).

258 Dental nomenclature follows Miller (1907), Freeman (1998), Garbino and Tavares (2018), and Garbino et al. (2020). Skull, dentition, and external characters were measured with digital calipers (to the nearest 0.01 mm). Total length, tail, hindfoot, ear, and body mass were recorded from skin labels and were only used for descriptive assessments (mean, range, and standard deviation).

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263 We log-transformed the variables and ensuring normality via Kolmogorov-Smirnov tests (Sokal and Rohlf 1995); then we performed two Analyses of Variances (ANOVA) with the 13
264 morphological variables to determine sex-based differences within subspecies. With no
265 significant differences found (all *p* < 0.05), we combined data from males and females for further
266 analysis. We used Principal Component Analysis (PCA) as a unified statistical approach to
267 investigate morphological variation in *Trachops cirrhosus*, applying it in two distinct contexts:
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270 (1) Morphological Variation Among Subspecies: To assess the morphometric divergence among
271 recognized *Trachops* subspecies, we first applied the PCA focusing on taxonomic identification
272 and current distribution ranges. This involved summarizing key characteristics of the dataset to
273 interpret complex relationships and trends in morphological traits (Sokal and Rohlf 1995).
274 Subsequently, PCA scores were subjected to a multivariate analysis of variance (MANOVA) and
275 Post-hoc multiple comparisons (Holm-Bonferroni correction) to detect significant morphometric
276 differences among subspecies (Rice 1989; Irschick and Shaffer 1997).

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278 (2) Geographic Variation Analysis: The second application of PCA was directed towards
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280 variable being a more specific geographic distribution area, rather than country boundaries.
281 Geographic areas were established as follows, based on Molinari et al. (2023):
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284 • CAm - Central America, from Mexico to Panama, delimited by the Atrato-San Juan Depression to the south, in Colombia.

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286 • WSAm - West South America, defined as the western slopes of the northern Andes, including the Colombian Western Cordillera, the Central Colombian Cordillera, and the Pacific coasts of Colombia, Ecuador, and Peru, to the Central Andes, delimited by the Atrato-San Juan Depression to the north and the Southern Andes to the south.
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290 • ESAm - East South America, demarcated as the upper forested slopes on the eastern side of the Andes and the low Amazon of Colombia, Ecuador, Peru, and Bolivia; the entire Amazon drainage of Brazil, including northeastern Bolivia, and the Orinoco drainages of Colombia and Venezuela.
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294 • NEC/NWV, NE Colombia and NW Venezuela, comprising the Orinoco llanos and Andean piedmont plus the Venezuelan Coast range, including the Guajira peninsula and the Maracaibo depression (Ferrer-Pérez et al. 2009).
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* **GS - the Guyana Shield**, including the Venezuelan states of Bolivar and Amazonas, and a portion of Delta Amacuro, the entire territories of Guyana, Surinam, and French Guyana, and parts of northern Brazil.

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* **AF - Atlantic Forest**, confined to the southernmost eastern region of Brazil, including the Mata Atlántica ecosystem.

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## Results

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**Specimens, tissue samples, and biorepositories:** Altogether, we measured 238 specimens of *Trachops cirrhosus* from all named subspecies (i.e., 202 *T. c. cirrhosus*, 31 *T. c. coffini*, and 5 *T. c. ehrhardti*), sourced from 7 scientific collections (Supplementary Table 1). Additionally, out of the 78 tissue samples, we successfully sequenced a total of 54 complete mitochondrial genomes (i.e., 47 *T. c. cirrhosus*; 5 *T. c. coffini*; 2 *T. c. ehrhardti*; Supplementary Table 1, Appendix 1). The remaining 24 individuals that could not be sequenced corresponded, mostly, to samples of ancient tissues preserved in formalin. 22 specimens were both measured and sequenced. To analyze geographic data, we used 173 unique collection locations, duly documented (Figure 2).

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**Phylogenetic analysis:** Monophyly of *Trachops* was recovered with strong support from both the ML analysis (bootstrap support [BS] = 100%), and from the Bayesian analysis (Bayesian posterior probability [BPP] = 1). There is broad agreement between analysis methods (ML and BI) for this dataset. We found two well-supported and genetically distant clades: one containing the subspecies *T. c. coffini*, nested within *T. c. cirrhosus* (Clade 1; Figure 3) while the other contained the subspecies *T. c. ehrhardti* (Clade 2, Figure 3).

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In Clade 1 we recovered 2 clear phylogroups: Phylogroup A comprising samples from Mexico, Guatemala, Honduras, Panama, western and northeastern Colombia, and northwestern Venezuela, and western Ecuador. According to the geographical groups analyzed in the

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morphometric analyzes (see Methodology and following section of Results), all these sequences correspond to the geographic groups CAm (Central America), WSAm (West South America) and NEC/NWV (NE Colombia and NW Venezuela), except for one sequence from Bolivia (FMNH 114877), which in the ML topology appears nested with individuals from western Ecuador. Within this phylogroup, there is a sequence from the province of El Petén, Guatemala, which is located 93 km away from the type locality of *T. c. coffini* (Guyo, Petén, Guatemala). The subspecies do not align with geographically distinct monophyletic groups, and furthermore, they do not match the geographical distributions historically associated with *T. c. cirrhosus* and *T. c. coffini* subspecies. Mean pairwise uncorrected sequence distances between what has been named as *T. c. cirrhosus* and *T. c. coffini* is 5.6% (Table 1), which is not different from the computed intraspecific divergence values of 5.5% in *T. cirrhosus* (Table 2).

We have identified a sister clade to Phylogroup A that we named as B (Figure 3), comprising individuals from Eastern Ecuador and Eastern Venezuela (Orinoco Basin), corresponding to the geographical group ESAm (East South America), as identified in our morphometric analyses. Besides these two strongly supported phylogroups, no further distinct phylogenetic relationships were found within Clade 1, although there is a noticeable clustering of samples from the northeastern region of South America, corresponding to the Guyana Shield.

According to the concordant phylogenies of ML and BI analyses, Clade 2 forms a monophyletic clade containing specimens from the southeastern region of São Paulo, in Brazil, which is geographically close to the state of Santa Catarina (type locality of *T. c. ehrhardti*). According to the geographical groups analyzed in the morphometric analyses (see Methodology and the following Results section), these sequences correspond to the geographic group AF (Atlantic Forest; Figure 3).

## 348 Morphologic and morphometric variation
Our examination of the *Trachops* subspecies highlights the morphological congruence, particularly in qualitative traits, between *T. c. cirrhosus* and *T. c. coffini*, contrasting with the distinct morphology of *T. c. ehrhardti*. Our scrutiny failed to discern marked morphological divergences between *T. c. cirrhosus* and *T. c. coffini*, challenging their current classification as distinct entities. The attributes previously considered distinctive, such as skull shape, rostrum morphology, dimensions of premolars and molars, and mandible features, displayed uniformity across these two subspecies in our geographical sampling (Figure 4). Table 3, illustrating variable ranges among the *Trachops* subspecies, suggests a considerable overlap, rendering these variables ineffective as discrete discriminators among the 3 subspecies.

In the Principal Component Analysis (PCA), two principal components cumulatively accounted for 73% of the total variance in our log-transformed dataset (Table 4). The PCA plot (Figure 5) reveals substantial overlap in the 2D morpho-space among the subspecies, attributed to their size and shape congruities. Notably, *T. c. cirrhosus* is positioned towards the right on PC1, indicating its relatively larger size compared to *T. c. coffini*, which is leaned towards the left. However, this positioning does not demarcate distinct groupings. PC1 predominantly shows size variations, influenced by skull length metrics such as Condyle-Incisive Length (CIL), Greatest Length of Skull (GLS), Maxillary Palatal Width (MPW), and Maxillary Toothrow Length (MTRL), each with factor loadings exceeding 0.8. Conversely, PC2 appears to reflect shape variations, primarily influenced by Palatal Breadth (PB).

Despite the apparent overlap in morpho-space depicted in Figure 5, our multivariate analysis of variance (MANOVA), employing Pillai's Trace test, identified statistically significant differences among the subspecies on both PC1 and PC2 (F = 16.373, *p* < 0.01). Further post-hoc multiple comparison analyses delineated a statistical distinction between *T. c. cirrhosus* and *T. c. coffini* in terms of size on PC1 (*p* < 0.01), but not between *T. c. cirrhosus* and *T. c. ehrhardti* (*p* > 0.05). Moreover, no statistical differences were noted between *T. c. coffini* and *T. c. ehrhardti* (*p* > 0.05).
These statistical disparities might be influenced by the limited sample size for *T. c. ehrhardti*, which does not sufficiently echo in morphological differentiation. The observed statistical differences, particularly between *T. c. cirrhosus* and *T. c. coffini*, are evident in the context of size, as indicated by PC1. However, the overarching morphological congruence, especially considering the overlap in the PCA plot, suggests that these subspecies, despite some statistical variance, share a degree of morphological commonality.

## Geographic variation

There is a noteworthy overlap in the 2D morpho-space between the geographic-based groups (Figure 6). The multivariate analysis of variance suggested there are differences between the geographic groups on PC1 and PC2, as suggested by Pillai's Trace test (F = 17.629, *P* < 0.01). Post-hoc multiple comparison analyses unveiled variation in size (PC1) among the samples. Specifically, there was a size difference between the samples from Central America (CAm) compared to those from East South America (ESAm), the Guyana Shield (GS), and North South America (NEC/NWV), but not when compared to the West South American (WSAm) samples. The plot illustrates that both CAm and WSAm samples cluster to the left along PC1, denoting their smaller size, but they do not end up forming a separate cluster. Conversely, the Atlantic Forest (AF) specimens were notably smaller in size compared to Guyana Shield (GS) samples, though not significantly different from the rest of the regions. In fact, the GS sample exhibited the largest size relative to all other groupings. Notably, NEC/NWV specimens did not display a clear grouping pattern. Regarding the PC2, AF specimens exhibited notable differences, particularly in variables such as palate width, in comparison to ESAm specimens. Additionally, a highly significant difference was observed between CAm and ESAm specimens.

Both qualitative and quantitative assessments reveal a noticeable degree of morphological and genetic similarity between the traditionally recognized subspecies *T. c. cirrhosus* and *T. c. coffini*. Moreover, our geographical analysis reveals that the distribution of morphological variation
among different regions does not align with the historical classification of subspecies. These findings strongly support their consideration as part of the same taxonomic group. Additionally, our investigation provides definitive evidence for elevating *T. c. ehrhardti* to the status of a distinct species, due to significant genetic and morphological disparities observed in specimens from the southeastern region of Brazil.

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## Discussion

411

Our investigation of *Trachops*, which integrates data from morphology, morphometrics, and molecular analysis encompassing almost the totality of its distribution range, has provided clarification into the taxonomic status of the species. Our main findings include the recognition of specific status of *T. ehrhardti*, and *T. c. coffini* as a junior synonym of *T. cirrhosus*. Incorporating *T. cirrhosus* specimens from Colombia, Ecuador, and Peru was crucial to our classification, as it represented the first taxonomic exploration of the species in the West South American region. We not only expanded the geographical scope of the analysis, but also shed light on previously unexplored genetic and morphological variations within *T. cirrhosus* populations.

420

### *Trachops* in phylogenetic context

422

The mitochondrial DNA (mtDNA) diversity in *Trachops cirrhosus* is significant, as evidenced by three previous studies. Using 10 partial sequences of the mitochondrial gene Cytb from localities such as Guatemala, Panama, French Guyana, and Brazil, *Ditchfield* (1996) identified 5 clades. These findings suggested that current subspecies classifications may underestimate haplotypic diversity, indicating that *Trachops* should be considered a complex. In this topology, *T. c. coffini* is not monophyletic as opposed to *T. c. cirrhosus*, and the author attributed this haplotypic diversity to reduced mobility or increased philopatry in *Trachops* across its geographic range, though these hypotheses remain unconfirmed (*Ditchfield* 1996).

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Later, Claire (2011) investigated pattern congruence between two independently evolving genetic regions (mitochondrial genome and Y chromosome), finding significant mitochondrial haplotypic divergence, but not in the Dby 7th intron region. Despite using more samples than Ditchfield (1996), the author did not include specimens from West South America. In this topology, *T. c. coffini* was once again not a monophyletic clade distinct from *T. c. cirrhosus*, lacking geographical congruence to support its separation as a subspecies.

Finally, Fonseca (2019) proposed a more integrative approach with a greater number of sequences using a dataset composed of 2341 bp of Cyt-b, COI, D-loop and STAT5A from 129 tissues. Samples from western South America, particularly Colombia, Ecuador, and Peru, were not included in this study. In consistency with Clare et al. (2011), the ML phylogeny did not render *T. c. coffini* as an independent clade of *T. c. cirrhosus*, with little genetic differentiation (Fonseca 2019). The author proposed that *Trachops* should be divided into two lineages recognized at the species level: *Trachops ehrhardti*, monotypic, and *T. cirrhosus*, with two subspecies (*T. c. cirrhosus* and *T. c. coffini*). However, the phylogenetic justification for maintaining *coffini* as a subspecies and its distribution was unclear.

In this study, we included an extensive taxon sampling with new localities from Colombia, Ecuador, and Peru, as well as information from complete mitochondrial genomes. Our findings revealed two reciprocally monophyletic clades, each exhibiting significant genetic divergence. The first clade, *T. cirrhosus*, notably includes sequences from northern Central America (*T. c. coffini*). This challenges the classical notion of *T. c. coffini* as a distinct subspecies, suggesting instead a more cohesive genetic identity within the *T. cirrhosus* clade. The second clade, *T. ehrhardti*, stands as a monophyletic group, sister to *T. cirrhosus*. These results prompt a reconsideration of *Trachops'* taxonomic structure, particularly the delineation of *T. ehrhardti* and *T. cirrhosus*, and the subspecies status of *T. c. coffini* within this framework.
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Trachops in the morphometric context

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When *T. coffini* was described as a separate species by Goldman (1925), he reported larger
measurements for *T. cirrhosus*. The author revised 18 specimens from the type locality of *T. coffini*
with others from Venezuela, Colombia, and Panama, assuming that they represented typical *T.
cirrhosus*, without specifying quantity or localities. However, his analysis, based on a limited
geographical range and a small sample size, raised questions about the robustness of these early
findings. Felten (1956a) and Felten (1956b) revisited this classification, incorporating a broader
geographic range, including Central America. His work, which suggested classifying the Central
American forms as *T. c. coffini* subspecies, highlighted the potential intraspecific variation within
the species. However, Felten's conclusions, particularly regarding the dental morphology and size
differences, were largely inferential, lacking quantitative rigor. The author proposed maintaining
the two subspecies and added *T. c. ehrhardti* as a subspecies, based on 3 specimens from
southeastern Brazil, with no apparent difference other than the smaller size, in relation to samples
from Colombia, recorded by Hershkovitz (1949), but similar in size to the samples from El
Salvador (Felten 1956a).

No other significant morphological study of *Trachops* was published until six decades later, when Fonseca (2019) revised specimens mostly from localities in Brazil. The quantitative analysis revealed a substantial overlap in the morphometric space across the three subspecies studied, notably between *T. c. cirrhosus* and *T. c. coffini*, challenging the notion of distinct morphometric differentiation between these subspecies. This finding was an indicative of a morphometric continuum rather than discrete categories. Qualitatively, she observed that *T. c. coffini*, exhibited a pronounced angle between the rostrum and the braincase in lateral view, a characteristic less evident in *T. c. cirrhosus*. In dental morphology, Fonseca (2019) noted that the first lower premolar (p1) in *T. c. coffini* possesses a unique shape, distinct to this subspecies, although the
specifics of this form were not delineated. Additionally, she reported that the m1 tooth in *T. c. coffini* demonstrates a more developed paraconid, compared to *T. c. cirrhosus*. The potential displacement of p3 to the labial surface in *T. c. coffini*, as opposed to the lingual surface in *T. c. cirrhosus*, suggests further morphological variability. Notably, Fonseca (2019) noted that the specimens from Costa Rica displayed morphological traits and cranial sizes more akin to *T. c. coffini*, diverging from the expected distribution patterns. Furthermore, the individuals from Panama exhibited traits from both *T. c. coffini* and *T. c. cirrhosus* in equal frequencies. This observation led Fonseca to propose a clinal variation in size across the species' range, with smaller individuals observed in Central America and larger ones in South America, positioning Panama as a zone of intergradation.

In our study, encompassing specimens from the western region of the Andes and northern South America, we did not observe the diagnostic characters reported by Fonseca (2019). Specifically, our samples from Central America did not exhibit the morphological differences between the northern and southern regions as previously proposed. This discrepancy, particularly regarding the distinct cranial angles in *T. c. coffini* and the unique shape of p1, suggests a less complex morphological variability within the *Trachops* genus. Even though the Principal Component Analysis (PCA) revealed a size variation on PC1, predominantly influenced by skull length metrics, the PCA plot depicted considerable overlap in morpho-space, indicating a high degree of morphological congruence. This overlap challenges the current subspecies classification based on size and shape traits. Contrary to the proposed latitudinal cline in size, our data indicate that the specimens from the western South America (WSAm), encompassing the Pacific regions of Colombia and Ecuador, and NEC/NWV are similar in size to those from Central America. This finding is consistent with zoogeographical studies that have identified a closer affinity between the bat fauna of the western Andes and Central America Koopman (1976); (1978); (1982); Hoffmann and Baker (2003); Clare (2011). Such patterns suggest the influence of historical events, ecological processes, and landscape heterogeneity (Manel et al. 2003) in shaping the
distribution and morphology of bat species, including *Trachops*. The role of heterogeneity of
habitats and environmental conditions as causes of morphological and genetic divergence
(Turmelle et al. 2011; Lindsey and Ammerman 2016) is also evident in our findings.

515

516 On the subspecific status of *Trachops cirrhosus coffini*

517

Despite previous assertions, notably by Fonseca (2019), our findings indicate that *T. c. cirrhosus*
and *T. c. coffini* lack distinct morphological synapomorphies and do not exhibit reciprocal
monophyly, as required for subspecies designation. This observation aligns with the criteria
outlined by Patten (2015), who mentions that a subspecies should be morphologically distinct
and geographically circumscribed, yet not necessarily forming a distinct genetic clade. According
to Molinari (2023), subspecies designation is appropriate for populations with significant and
heritable morphological differences, even if genetic differentiation is insufficient for species-level
recognition. Our analyses, however, reveal that the morphological distinctions traditionally used
to separate *T. c. coffini* and *T. c. cirrhosus* do not meet these standards. The lack of clear
geographical circumscription, combined with our findings that these groups are not
morphologically diagnosable nor genetically distinct, challenges the validity of their current
subspecific status.

530

The reliance on size-only differences is contentious. Molinari (2023) underscores that diagnostic
morphological characters should not be attributed to phenotypic plasticity. Considering studies
indicating size variation as a source of morphological plasticity in bats (McLellan 1984; Jarrín-V
et al. 2010; Jarrín-V. & Menéndez-Guerrero 2011; López-Aguirre et al. 2015), the use of body size
as a sole criterion for taxonomic decisions becomes problematic. Geographic variation in body
size, influenced by a multitude of factors, including genetic adaptations and environmental
conditions affecting growth rates, further complicates its reliability as a distinguishing feature
(McLellan 1984; Berry et al. 1987; Ebenhard 1990; Jarrín-V et al. 2010; Jarrín-V. & Menéndez-
539 Guerrero 2011; López-Aguirre et al. 2015). Additionally, methodological inconsistencies, including instrumental and human errors, can lead to data discrepancies (Fox et al. 2020), casting doubt on the robustness of size-based differentiation. Our study, therefore, suggests that the current classification of *T. c. coffini* as a subspecies of *Trachops cirrhosus* lacks support, both morphologically and genetically.

544

545 **On the specific status of *T. ehrhardti***

546

547 *Trachops ehrhardti* should be elevated to the species status based on the presence of clear diagnosable morphological characters, geographically circumscription in the Atlantic forests of South Brazil (Mata Atlántica), and substantial molecular divergence. Broader cranial measurements (BB, MPW, ZB) allow *T. cirrhosus* to be separated from *T. ehrhardti* and both species form reciprocally monophyletic clades separated by a genetic divergence of over 8%. It is its geographic isolation that would primarily imply loss of ecological exchangeability, and possibly mating impediments with *Trachops cirrhosus*. Originally proposed by Haffer (1969), the Refuge Theory for the Neotropics suggests that the Pleistocene glaciation cycles created contraction and subsequent expansion of forested areas that, in turn, would create allopatry between populations of the same forest-dwelling species, leading to intraspecific differentiation and consequently speciation. Paleopalynological research indicates the high dynamism of Neotropical forested regions: the Atlantic Forest and the Amazon were connected in the past (Vivo 1997), separating themselves as increasing aridity in the Tertiary triggered the formation of the belt of xeromorphic formations between them (Martins et al. 2009). There seemed to be a predominantly arboreal vegetation during most of the Pleistocene, with typical Amazonia and Atlantic Forest tree species found in what it is now the dry diagonal that separates these two biomes. The extent to which these climatic fluctuations and associated vegetation changes affected the patterns of distribution and diversification of the fauna remains a central question in understanding the evolution of forest-associated taxa.
566

567 Implications to conservation

568

569 Recognized bat diversity has increased due to new species descriptions and taxa raised from
570 subspecific level or synonymy (Burgin et al. 2018), but also as a result of clarification of cryptic
571 species within several genera (e.g. Platyrrhinus (Velazco et al. 2023); Glossophaga (Calahorra-
572 Oliart et al. 2021) or Sturnira (Yánez-Fernández et al. 2023). These changes in taxonomy may have
573 an impact on the conservation of species. The genus *Trachops* is composed of two monotypic
574 species: *Trachops cirrhosus* maintains its “Least Concern” conservation status at the global level,
575 with no major threats known throughout its range (Miller et al. 2015). However, it is important to
576 revise the status of *Trachops ehrhardti*. The current range of *T. ehrhardti* is restricted to
577 southeastern Brazil. The Atlantic Forest is one of the world's leading biodiversity 'hotspots'; these
578 areas possess the highest concentrations of species endemism, and simultaneously, are the most
579 severely threatened by habitat loss (Baptista & Rudel 2006). In this biome, the persistent land
580 cover is made up of 476 000 Km², which comprises 9% of the Brazil's land area (Souza et al. 2020),
581 but is highly fragmented by roads and urban centers, and immersed in a large agriculture matrix,
582 resulting in less than 12% of old secondary forest cover (i.e., >30 years) (de Rezende et al. 2015;
583 Souza et al. 2020). If this scenario remains unchanged, then elevating *Trachops ehrhardti* to a
584 species should trigger immediate protection efforts in conservation.

585

586 We present a revised description of the genus *Trachops* and the subspecies *Trachops cirrhosus* and
587 *Trachops ehrhardti* below.

588

589 Taxonomy

590 The genus *Trachops* can be easily recognized by their highly specialized warty outgrowths or
591 protuberances on the lips and chin. The margin of the nose leaf is finely serrated on the edge,
592 anteriorly connected to the upper lip (Spix 1823). Ears are large and anteriorly covered with hairs
593 projecting beyond anterior margins (Goldman 1925); folds inside the pinnae are well marked. The
tail is short and appears on the upper side of a broad interfemoral membrane (Goodwin and
594
595 Greenhall 1961). The ventral fur is characterized by a pale brown coloration (Spix 1823). Dorsal
596 fur is cinnamon-brown, varying to darker shades in some specimens (Goodwin and Greenhall
597
598
599

**Trachops cirrhosus** Gray, 1847

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*Vampyrus cirrhosus* Spix, 1823:64. No type locality is stated in Spix's description, but on
page 53 Spix said the bats were collected in Brazil. Type locality restricted to Para, Brazil,
by Husson (1962: 115).

*Phyllostoma*. *cirrhosus*: Fischer, 1829:126. Name combination.

*Vampyris cirrhosum*: Gray, 1847:481. Emendation of *Vampyrus cirrhosus* Spix

*Trachops fuliginosus*: Gray, 1865:14. Type locality "Pernambuco," Brazil (= *Vampyrus cirrhosus* Spix).

*Phyllostoma angusticeps* Gervais, 1856: 47. Type locality "province de Bahia, au Brasil."

*Tylostoma mexicana* Saussure, 1860 :484. Type locality "les régions chaudes du Mexique."

*Trachyops cirrhosus*: Dobson, 1878:481. First use of name combination and incorrect
subsequent spelling of *Trachops* Gray, 1847.

*Trachops coffini* Goldman, 1925:23. Type locality "Guyo, Petén, Guatemala." Type locality
restricted to "El Gallo, 8 mi. west Yaxhá, on the Remate-El Cayo trail, Petén, Guatemala" by
de la Torre (1956:189).

*Trachops cirrhosus coffini*: Felten, 1956a:189. Type locality restricted to Para, Brazil, by
Husson (1962: 115).

This species is monotypic.
619 **Amended distribution and habitat:** This species is widely distributed in eastern and southern Mexico, extending from Veracruz, in the southeast, southeastward through Central America, into South America, including Colombia, Venezuela, the Guyanas, Ecuador, northern and central Peru, Bolivia, Trinidad, as well as northern, southern, and central Brazil (Figure 7). This species occurs at elevations from sea level (e.g., Belem, Para, Brazil) up to 1800 m (Las Tolas, Pichincha, Ecuador; (Arcos et al. 2007). Records of *Trachops cirrhosus* are predominantly associated with habitats in humid tropical forests, as well as sub-montane and montane forests.

626

627 **Emended diagnosis and comparison:** *Trachops cirrhosus* is a relatively robust (Spix 1823), medium-size bat (FA: 49.57-64.70 mm; GLS 24.70-31.24 mm). Overall size is larger than *Trachops ehrhardti* (Table 5). The skull is large and elongated. The elevated braincase above the rostrum was previously believed to be a distinguishing characteristic, potentially separating the subspecies *cirrhosus* and *coffini*. The latter was described as having a smaller and slenderer shape compared to *T. c. cirrhosus*, although similar to *T. c. ehrhardti*. This characteristic is not diagnostic; rather, it displays variability across the entire species distribution. The sagittal crest may or may not be developed in males and females, contrary to the well-developed sagittal crest character mentioned by Goodwin and Greenhall (1961). Poorly developed sagittal crest is common in most specimens reviewed for the entire area of distribution. However, the faint notch in the cutting edge of the upper incisors (Goldman 1925), is variable. Inferior premolar 1 (p1) is taller compared to the same tooth in *T. c. ehrhardti* which is wider overall. The first lower premolar is three-fourths the height of the third premolar but wider in cross-section (Dobson 1878). The mandible of *T. cirrhosus* is broad and robust; upper and lower premolars are relatively narrower than in *T. ehrhardti*. The dentary is tall and has a contracted premolar toothrow, and an expanded molar toothrow, with the last molar being closer to the fulcrum, a high coronoid process, and an expanded angular process (Figure 4).

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644
645 **Remarks:** The character of m1 with a less developed paraconid in *T. c. cirrhosus* than in *T. coffini*
646 mentioned by Dobson (1878) was not evident as a diagnostic character.

647
648 **Natural history:** *Trachops cirrhosus* is a large-eared gleaning bat that hunts by listening for prey-
649 generated sounds (Obrist et al. 1993). It roosts in caves, hollow trees, road culverts, sewer
650 systems, and buildings, in groups of up to 50 individuals (Hall and Dalquest 1963; Fleming et al.
651 1972; Alvarez-Castañeda and Álvarez 1991; Kalko et al. 1999; Halczok et al. 2018). The relatively
652 sedentary foraging behavior of these gleaners is reflected in their wing morphology, a
653 characteristic that influences their use of small foraging areas and short commuting distances.
654
655 They have been observed to engage in two distinct flight patterns while foraging: long flights that
656 extend for several uninterrupted minutes and short sally flights lasting less than one minute
657 (Cramer et al. 2001). These bats possess relatively short and broad wings, an adaptation that
658 enhances their maneuverability, particularly in environments filled with obstacles (Marinello &
659 Bernard 2014). However, this wing morphology comes with a trade-off, as it makes continuous
660 flight over extended distances energetically costly (Norberg & Rayner 1987; Kalko et al. 1999).

660 *Trachops cirrhosus* hunts for prey in continuous flight, presumably depending on prey availability.
661
662 Prey is usually taken from the substrate (gleaning mode) in a brief landing or may be caught
663 occasionally on the wing (aerial mode) (Kalko et al. 1999). *Trachops cirrhosus* is a carnivorous bat
664 that feeds on a wide variety of prey species, including insects, frogs, lizards, and other small
665 vertebrates (Gardner 1977; Pine and Anderson 1979; Barclay et al. 1981; Tuttle & Ryan 1981;
666 Bonato and Facure 2000; Bonato et al. 2004; Giannini and Kalko 2005; Page and Jones 2016).
667
668 Additionally, it has been observed consuming fruits and seeds (Whitaker & Findley 1980;
669 Humphrey et al. 1983; Cramer et al. 2001). Foraging areas of 3 to 12 ha and commuting distances
670 between roost and foraging areas are < 2 km (Kalko et al. 1999). The emergence time and activity
671 peak of *T. cirrhosus* coincided with the maximum calling activity of the Tungara frogs,
Leptodactylidae, (Tuttle & Ryan 1981; Ryan et al. 1983). The number and duration of long flights
reduce as the calling declines; giving way to an increased frequency of short flights, which might
also indicate a switch from frogs to other prey such as insects (Belwood 1990; Kalko et al. 1996, 1999). This bat is very flexible in its responses to prey calls by updating acoustic information with echo acoustic and gustatory cues, as it approaches potential prey, enabling bats to avoid potentially lethal mistakes (Page and Jones 2016). Captive studies show that cues can also be learned socially and transmitted across individuals (Jones et al. 2013; Page and Jones 2016; Flores et al. 2020).

Very little is known about the reproductive biology of *T. c. cirrhosus*. Females give birth to one offspring at a time coinciding with the start of the rainy season (Flores and Page 2017), but the gestation period length is unknown. Females of this species have been reported to be pregnant in April to March and December (Villa-R 1967; Álvarez-Castañeda and Álvarez 1991). In Brazil, Trajano (1984) suggested a polyestrous reproductive pattern, with two annual birth peaks, one before and the other after August. In Trinidad and Tobago, Goodwin and Greenhall (1961) found a colony of *T. cirrhosus* composed of 6 individuals of both sexes, including pregnant females in March (Bredt et al. 1999). The social structure of *T. cirrhosus* is still not fully understood, but a recent six-year study in Panama showed evidence of female philopatry and preferred co-roosting associations in both sexes; kin-biased associations were also detected among pairs of females but not males (Flores et al. 2020).

During the mating season, reproductive males have enlarged testes and create an odorous substance that is smeared on their forearm, called forearm crust. Flores and Page (2017) and Flores et al. (2019) discovered that fringe-lipped males scratch their body with one hind claw, including the area around a prominent mid-ventral chest gland, insert the same hind claw into the mouth, and then repeatedly lick the forearm. Apparently, this substance is not related to female preferences since two-thirds of females selected the scent of a male without forearm crust, but it would play a fundamental role in male-male interactions (Flores et al. 2019).
699 Trachops ehrhardti Felten, 1956

700
701 Trachops cirrhosus ehrhardti Felten, 1956b:369. Type locality "Joinville, Sta. Catarina,
702 Brazil."

703
704 Distribution and habitat: Southeastern Brazil. Specimens are known from the humid
705 environments of the Atlantic Forest of the states of Santa Catarina, Parana, Sao Paolo, Mina Gerais,
706 Rio de Janeiro and Espiritu Santo (Figure 7).

707
708 **Diagnosis:** Medium-size bat (FA: 54-60 mm; GLS 26.90-27.70 mm). Smaller than *Trachops cirrhosus* (Table 5). The ventral fur from the base of the hair shows a lighter shade of brown that
709 gradually transitions to an ashy color at the tips. Dorsal fur is characterized by a reddish-brown
710 to cinnamon-brown hue. The base of the hair appears whitish, creating a contrast with the slightly
711 ashy tone at the tips. Underparts are light brownish with a grayish tint. This coloration is
712 attributed to the presence of light-colored tips on the hairs, and there are white-tipped hairs
713 specifically found on the animal's underparts. The pinnae are hairy, with marked folds. Lips and
714 chin are ornamented with wart-like protuberances and ears are large and clothed with hairs
715 projecting conspicuously beyond anterior margins as in *Trachops cirrhosus* (Goldman 1925).
716
717 Skull is noticeably smaller in comparison to *T. cirrhosus*. Braincase is smaller, forming an angle
718 with a less pronounced rostrum than in *T. cirrhosus*, however, this character shows more degree
719 of variation. Sagittal crest is not developed. In the analyzed skulls, which belong to the type series,
720 it is observed that the lingual edges of the w-shaped stylar shelf are longer. The molar crowns are
721 relatively longer, and in the dorsal view, the parastyle, mesostyle, and metastyle of the second and
722 third upper molars are further away from the maxillary bone. The formed cusps are further apart
723 than in *T. cirrhosus*, giving them a broader appearance. These characters could be important to
724 identify individuals from the eastern areas of Brazil where there could be contact between *T.
725 cirrhosus* and *T. ehrhardti* (Figure 4).
726

727

**Natural history:** To the best of our knowledge, precise data on the natural history of this population is currently lacking, although it is expected to share similarities with *T. cirrhosus*. This species inhabits the Atlantic Forest, which encompasses both lowland and montane systems along the Atlantic coast of southeastern Brazil, as well as the contiguous moist-subtropical forest of the Parana basin (Pavan et al. 2016). Similar to *T. cirrhosus*, it is likely that they seek refuge in caves, hollow trees, culverts, and buildings, often in small groups comprising a few tens of individuals. The species might prey on insects, frogs, lizards, and other small vertebrates. Currently, no information is available regarding their reproductive behavior.

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734

735

736

**Remarks:** The species was described by Felten (1956b) based on 3 specimens collected in 1908 by W. Ehrhardt at Joinvile in the Brazilian state of Santa Catarina, considerably expanding the southern limit of distribution of the species in South America. The only distinctive characteristic of the subspecies mentioned by Felten (1956b) is its size, notably smaller in relation to animals from the northern part of South America and similar in size to the former subspecies *T. c. coffini*. In 2019, in a degree dissertation from the Federal University of Espírito Santo (see Fonseca, unpublished data), it was proposed, based on an integrative approach to morphological, ecological, and phylogenetic evidence, that *T. cirrhosus ehrhardti* should be elevated to the species category; however, the species has not been formally described nor has the taxonomy been validated.

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746

747

**Comparisons:** The size and shape of the skull of the two species is similar, but in *Trachops ehrhardti* it is smaller on average. In *T. cirrhosus*, the skull is larger and more elongated than in *T. ehrhardti*, and the braincase is more elevated above the rostrum. However, this character shows a degree of variation. *T. cirrhosus* presents a faint notch in the cutting edge of the upper incisors while is broad, with an open groove leading to a distinct notch in the cutting edge in *T. ehrhardti* (Goldman 1925), but this characteristic is also variable. The first lower premolar 1 (p1) in *T*.

748

749

750

751

752
753 *ehrhardti* is wider overall compared to the same teeth in *T. cirrhosus* which is taller than wider.

754 Molariforms tooththrow in *T. cirrhosus* is longer than in *T. ehrhardti*, *Trachops cirrhosus* mandible is broader and more robust than in *T. ehrhardti*. In *T. cirrhosus*, the dentary is tall and has a contracted premolar tooththrow, and an expanded molar tooththrow, with the last molar being closer to the fulcrum, a high coronoid process, and an expanded angular process. This could be associated with a higher bite force (Nogueira et al. 2009). Regarding the width of the cranium, 3 measurements are broader in *cirrhosus* than *ehrhardti* (i.e. Breadth of brain case, Mastoid process breadth, and Zygomatic breadth).

761

762

763 Acknowledgments

764 We extend our gratitude to the FSPI-Doctoral Schools Project of the French Embassy in Ecuador, supported by the Ministry of Europe and Foreign Affairs. This work has greatly benefited from the "Investissement d'Avenir" grants overseen by the Agence Nationale de la Recherche (CEBA, ANR-10-LABX-25-01; TULIP, ANR-10-LABX-0041). We are thankful to Marisa Surovy (American Museum of Natural History), Darrin Lunde (National Museum of Natural History), Nicolás Reyes-Amaya (Instituto de Investigación de Recursos Biológicos Alexander von Humboldt), Oscar E. Murillo-García (Universidad del Valle), and Edith Montalvo (Museo de Historia Natural Gustavo Orcés-V.) for providing access to invaluable specimens. We would also like to thank Adam Ferguson and Bruce Paterson from the Field Museum of Natural History, Marie L. Campbell and Joseph Cook from the Museum of Southwestern Biology, Jacqueline Miller and Burton Lim from the Royal Ontario Museum, and Dr. Irina Ruf from the Senckenberg Naturmuseum Frankfurt for their generosity in providing tissue samples. For their work on the mtDNA sequencing and assembly, we express our profound thanks to Alexandra Bialonski and Marike Petersen from the Bernhard Nocht Institute for Tropical Medicine. We also want to acknowledge Ruben D. Jarrín from Pontificia Universidad Católica del Ecuador and Anika Vogel from the Senckenberg Research Institute for their photographic contributions. Santiago F. Burneo deserves a special
780 mention for granting access to specimens under his care at QCAZ and his help with the maps

781 presented on this manuscript. Our sincere appreciation goes to Diego Tirira and the anonymous
782 reviewers whose insights and comments have significantly enhanced this manuscript.

783

784 **References**

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1189 https://www.researchgate.net/publication/322953093_Guia_de_campo_de_los_mamiferos_del_Ecuador

1191
1192 Trajano E. 1984. Ecologia de populações de morcegos cavernícolas em uma região cárstica do
1193 sudeste do Brasil. *Revista Brasileira de Zoologia* 2: 255-320. https://doi.org/10.1590/S0101-81751984000100001

1195
1196 Turmelle A.S., Kunz T.H., Sorenson M.D. 2011. A tale of two genomes: contrasting patterns of
1197 phylogeographic structure in a widely distributed bat. *Molecular Ecology* 20: 357-375.
1198 https://doi.org/10.1111/j.1365-294x.2010.04947.x

1199
1200 Tuttle M.D., Ryan M.J. 1981. Bat predation and the evolution of frog vocalizations in the neotropics.
1201 *Science* 214: 677-678. https://doi.org/10.1126/science.214.4521.677

1202
1203 Velazco P.M. 2005. Morphological Phylogeny of the Bat Genus *Platyrrhinus* Saussure, 1860
1204 (Chiroptera: Phyllostomidae) with the Description of Four New Species. *Fieldiana Zoology* 2005:
1205 1-53. https://doi.org/10.3158/0015-0754(2005)105[1:MPOTBG]2.0.CO;2

1206
1207 Velazco P.M., Ly G., McAllister J., Esquivel D.A. 2023. Geographic variation in select species of the
1208 bat genus *Platyrrhinus*. *Therya* 14: 121-130. https://doi.org/10.12933/therya-23-2208

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1218
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1221
1222 Williams S.L., Genoways H.H. 2008. Subfamily Phyllostominae Gray, 1825. Mammals of South America. Volume 1: Marsupials, Xenarthrans, Shrews, and Bats. Gardner AL, editor. Chicago
1223
1224 (IL,USA): The University of Chicago Press.
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1226
1227 Yánez-Fernández V., Marchán-Rivadeneira M.R., Velazco P.M., Burneo S.F., Tinoco N., Camacho
1228 M.A. 2023. On the Taxonomic Identity of *Sturnira* nana Gardner and O'Neil, 1971 (Chiroptera:
1229 Phyllostomidae), from Ecuador, with the Description of a New Species of *Sturnira*. *American
1230 Museum Novitates* 2023: 1-28. https://doi.org/10.1206/4001.1
1231 TABLES

1232

1233

1234

1235

1236

**Table 1.** Estimates of evolutionary divergence across sequence pairs between groups are presented. The number of base differences per site, calculated by averaging over all sequence pairs between groups, is depicted below the diagonal and scaled as percentages.

<table><thead><tr><th></th><th>T. c. cirrhosus</th><th>T. c. coffini</th><th>T. c. ehrhardti</th><th>Outgroup</th></tr></thead><tbody><tr><td>T. c. cirrhosus</td><td></td><td>0.0013</td><td>0.0018</td><td>0.0023</td></tr><tr><td>T. c. coffini</td><td>5.6%</td><td></td><td>0.0022</td><td>0.0025</td></tr><tr><td>T. c. ehrhardti</td><td>8.2%</td><td>8.6%</td><td></td><td>0.0025</td></tr><tr><td>Outgroup</td><td>17.7%</td><td>17.8%</td><td>17.8%</td><td></td></tr></tbody></table>

1237

1238
1239 **Table 2.** Estimates of evolutionary divergence across sequence pairs within groups
1240 Standard error estimate(s) are shown in the second column and were obtained by a bootstrap
1241 procedure (1000 replicates).

1242

<table><thead><tr><th></th><th>Distance %</th><th>S.E. (%)</th></tr></thead><tbody><tr><td><i>T. c. cirrhosus</i></td><td>5.5</td><td>0.0</td></tr><tr><td><i>T. c. coffini</i></td><td>1.9</td><td>0.0</td></tr><tr><td><i>T. c. ehrhardti</i></td><td>0.0</td><td>0.0</td></tr><tr><td>Outgroup</td><td>0.1</td><td>0.0</td></tr></tbody></table>

1243
<table><caption>**Table 3.** Measurements (mm) and body masses (g) of *Trachops c. cirrhosus*, *T. c. coffini* and *T. c. ehrhardti*. Descriptive measures: minimum-maximum; arithmetic mean (standard deviation).</caption><thead><tr><th>Variable</th><th>T. cirrhosus<br>cirrhosus<br>(N = 202)</th><th>T. cirrhosus coffini<br>(N = 31)</th><th>T. cirrhosus<br>ehrhdarti<br>(N = 5)</th></tr></thead><tbody><tr><td>Breadth of brain case (BB)</td><td>11.07-12.97; 11.60<br>(0.28)</td><td>10.33-12.03; 11.49<br>(0.26)</td><td>11.30-11.50; 11.40<br>(0.10)</td></tr><tr><td>Palatal width at canines (C-C)</td><td>5.72-6.85; 6.22<br>(0.21)</td><td>5.26-6.68; 5.98<br>(0.26)</td><td>5.4-5.80; 5.63 (0.21)</td></tr><tr><td>Condylocanine length (CCL)</td><td>22.23-26.79; 25.01<br>(0.71)</td><td>20.93-26.12; 24.17<br>(0.86)</td><td></td></tr><tr><td>Condyloincisive length (CIL)</td><td>23.30-27.95; 26.03<br>(0.69)</td><td>21.87-26.78; 25.10<br>(0.79)</td><td>23.90-24.50; 24.20<br>(0.30)</td></tr><tr><td>Coronoid height (COH)</td><td>4.97-6.80; 5.85<br>(0.30)</td><td>4.79-6.28; 5.42<br>(0.31)</td><td></td></tr><tr><td>Dentary length (DENL)</td><td>17.41-20.48; 18.97<br>(0.60)</td><td>16.11-19.93; 18.15<br>(0.67)</td><td></td></tr><tr><td>Greatest length of skull (GLS)</td><td>26.13-31.24; 29.21<br>(0.79)</td><td>24.70-30.02; 28.27<br>(0.88)</td><td>26.90-27.70; 27.20<br>(0.44)</td></tr><tr><td>Breadth across upper molars<br>(M2-M2)</td><td>9.15-11.03; 10.14<br>(0.32)</td><td>8.34-10.86; 9.77<br>(0.38)</td><td>9.80</td></tr><tr><td>Mandibular tooththrow length<br>(MANDL)</td><td>10.71-12.40; 11.42<br>(0.34)</td><td>7.61-11.78; 10.92<br>(0.54)</td><td></td></tr><tr><td>Molariform tooththrow<br>(MLTRL)</td><td>6.68-7.81; 7.22<br>(0.21)</td><td>6.07-7.67; 6.96<br>(0.26)</td><td></td></tr><tr><td>Mastoid (process) breadth<br>(MPW)</td><td>12.78-14.96; 13.70<br>(0.41)</td><td>11.40-14.31; 13.21<br>(0.46)</td><td>13-13.40; 13.20<br>(0.20)</td></tr><tr><td>Maxillary tooththrow (MTRL)</td><td>9.50-11.33; 10.51<br>(0.32)</td><td>8.57-10.93; 10.11<br>(0.40)</td><td>9.60-10; 9.8 (0.20)</td></tr><tr><td>Post orbital constriction<br>breadth (PB)</td><td>4.85-5.76; 5.34<br>(0.17)</td><td>4.66-5.71; 5.11<br>(0.18)</td><td>5</td></tr><tr><td>Palatal length (PL)</td><td>10.12-12.72; 11.40<br>(0.47)</td><td>8.92-12.42; 11.00<br>(0.52)</td><td></td></tr><tr><td>Zygomatic breadth (ZB)</td><td>12.46-15.97; 14.53<br>(0.48)</td><td>12.06-15.70; 13.96<br>(0.58)</td><td>13.3-14.10; 13.77<br>(0.42)</td></tr><tr><td>Braincase height (BCH)</td><td>9.24-12.20; 10.84<br>(0.47)</td><td>9.39-12.25; 10.63<br>(0.42)</td><td></td></tr><tr><td>Calcar length (CL)</td><td>9.19-18.21; 13.29<br>(1.52)</td><td>10.84-16.55; 13.64<br>(1.30)</td><td>11.62</td></tr><tr><td>Forearm length (FA)</td><td>53.36-64.70; 59.23<br>(2.25)</td><td>49.57-62.11; 57.88<br>(2.51)</td><td>53.99-60; 57.29<br>(2.43)</td></tr><tr><td>Hindfoot length (HF)</td><td>12.40-23; 17.38<br>(2.35)</td><td>13.92-22; 17.66<br>(2.04)</td><td>11.47-16.01; 13.74<br>(3.21)</td></tr><tr><td>Metacarpal III (MET-III)</td><td>35.31-52.88; 48.18<br>(2.49)</td><td>37.56-53.40; 47.63<br>(2.27)</td><td>39.45-52.30; 47.56<br>(4.91)</td></tr><tr><td>Metacarpal IV (MET-IV)</td><td>37.12-55.48; 50.24<br>(2.56)</td><td>37.98-54.43; 49.34<br>(2.36)</td><td>40.57-53.30; 49.51<br>(5.18)</td></tr><tr><td>Metacarpal V (MET-V)</td><td>39.04-58.86; 53.53<br>(2.73)</td><td>41.15-58.13; 53.25<br>(2.20)</td><td>42.85-55.60; 51.79<br>(5.2)</td></tr><tr><td>Tibia length (TiL)</td><td>16.82-29.69; 25.90<br>(1.84)</td><td>20.38-28.90; 24.94<br>(1.45)</td><td>20.61-25.16; 23.09<br>(1.66)</td></tr><tr><td>Tail length (T)</td><td>10-28; 16.93 (3.25)</td><td>10-22; 16.11 (2.75)</td><td>14.38</td></tr><tr><td>Total length (TL)</td><td>59.70-128; 96.16<br>(11.35)</td><td>56.25-107; 91.09<br>(15.94)</td><td></td></tr><tr><td>Ear length (E)</td><td>23.07-39; 31.10</td><td>20-39; 31 (4.06)</td><td>27.65-31; 29.33</td></tr></tbody></table>
**Weight (W)** (4.06) (2.37)
16-55; 34.48 (7.74) 22-46; 33.19 (6.99)
1248 **Table 4.** Loadings, eigenvalues and percentage of variance for 2 principal components

1249 from a PCA of the 13 linear measurements for adult specimens of *Trachops*.

<table><thead><tr><th>Variables</th><th>PC1</th><th>PC2</th></tr></thead><tbody><tr><td>Breadth of brain case (BB)</td><td>.554</td><td>.280</td></tr><tr><td>Palatal width at canines (C-C)</td><td>.778</td><td>.314</td></tr><tr><td>Condyloincisive length (CIL)</td><td>.919</td><td>.068</td></tr><tr><td>Greatest length of skull (GLS)</td><td>.919</td><td>.071</td></tr><tr><td>Breadth across upper molars (M2-M2)</td><td>.756</td><td>.293</td></tr><tr><td>Mastoid (process) breadth (MPW)</td><td>.820</td><td>.212</td></tr><tr><td>Molariform toothrow (MLTRL)</td><td>.872</td><td>.067</td></tr><tr><td>Post orbital constriction breadth (PB)</td><td>.383</td><td>.698</td></tr><tr><td>Zygomatic breadth (ZB)</td><td>.864</td><td>.128</td></tr><tr><td>Forearm length (FA)</td><td>.743</td><td>-.180</td></tr><tr><td>Metacarpal III (MET-III)</td><td>.690</td><td>-.555</td></tr><tr><td>Metacarpal IV (MET-IV)</td><td>.800</td><td>-.515</td></tr><tr><td>Metacarpal V (MET-V)</td><td>.786</td><td>-.546</td></tr><tr><td>Eigenvalues</td><td>7.787</td><td>1.731</td></tr><tr><td>% of explained variance</td><td>59.9</td><td>13.3</td></tr><tr><td>% of cumulative variance</td><td>59.9</td><td>73.2</td></tr></tbody></table>

1250
1251 **Table 5.** Measurements (mm) and body masses (g) of *Trachops cirrhosus* and *T. ehrhardti*.

1252 Descriptive measures: minimum-maximum; arithmetic mean (standard deviation)

<table><thead><tr><th>Variable</th><th>T. cirrhosus<br>N = 233</th><th>T. ehrhdarti<br>N = 5</th></tr></thead><tbody><tr><td>Breadth of brain case (BB)</td><td>10.33-12.97; 11.56 (0.28)</td><td>11.30-11.50; 11.40 (0.10)</td></tr><tr><td>Palatal width at canines (C-C)</td><td>5.26-6.85; 6.13 (0.26)</td><td>5.40-5.80; 5.63 (0.21)</td></tr><tr><td>Condylocanine length (CCL)</td><td>20.93-26.79; 24.70 (0.87)</td><td></td></tr><tr><td>Condyloincisive length (CIL)</td><td>21.87-27.95; 25.67 (0.85)</td><td>23.90-24.50; 24.20 (0.30)</td></tr><tr><td>Coronoid height (COH)</td><td>4.79-6.80; 5.71 (0.36)</td><td></td></tr><tr><td>Dentary length (DENL)</td><td>16.11-20.48; 18.70 (0.72)</td><td></td></tr><tr><td>Greatest length of skull (GLS)</td><td>24.70-31.24; 28.86 (0.94)</td><td>26.90-27.70; 27.20 (0.44)</td></tr><tr><td>Breadth across upper molars (M2-M2)</td><td>8.34-11.03; 10 (0.39)</td><td>9.80</td></tr><tr><td>Mandibular toothrow length (MANDL)</td><td>10.06-12.40; 11.05 (0.40)</td><td></td></tr><tr><td>Molariform toothrow (MLTRL)</td><td>6.07-7.81; 7.12 (0.27)</td><td></td></tr><tr><td>Mastoid (process) breadth (MPW)</td><td>11.40-14.96; 13.51 (0.49)</td><td>13-13.40; 13.20 (0.20)</td></tr><tr><td>Maxillary toothrow (MTRL)</td><td>8.57-11.33; 10.36 (0.40)</td><td>9.60-10; 9.8 (0.20)</td></tr><tr><td>Post orbital constriction breadth (PB)</td><td>4.66-5.76; 5.25 (0.21)</td><td>5</td></tr><tr><td>Palatal length (PL)</td><td>8.92-12.72; 11.25 (0.53)</td><td></td></tr><tr><td>Zygomatic breadth (ZB)</td><td>12.06-15.97; 14.31 (0.59)</td><td>13.3-14.10; 13.77 (0.42)</td></tr><tr><td>Braincase height (BCH)</td><td>9.24-12.25; 10.76 (0.46)</td><td></td></tr><tr><td>Calcar length (CL)</td><td>9.19-18.21; 13.41 (1.44)</td><td>11.62</td></tr><tr><td>Forearm length (FA)</td><td>49.57-64.70; 58.71 (2.43)</td><td>53.99-60; 57.29 (2.43)</td></tr><tr><td>Hindfoot length (HF)</td><td>12.40-23; 17.49 (2.24)</td><td>11.47-16.01; 13.74 (3.21)</td></tr><tr><td>Metacarpal III (MET-III)</td><td>35.31-53.40; 47.93 (2.41)</td><td>39.45-52.30; 47.56 (4.91)</td></tr><tr><td>Metacarpal IV (MET-IV)</td><td>37.12-55.48; 49.93 (2.51)</td><td>40.57-53.30; 49.51 (5.18)</td></tr><tr><td>Metacarpal V (MET-V)</td><td>39.04-58.86; 53.42 (2.54)</td><td>42.85-55.60; 51.79 (5.2)</td></tr><tr><td>Tibia length (TiL)</td><td>16.82-29.69; 25.57 (1.76)</td><td>20.61-25.16; 23.09 (1.66)</td></tr><tr><td>Tail length (T)</td><td>10-28; 16.61 (3.08)</td><td>14.38</td></tr><tr><td>Total length (TL)</td><td>56.25-128; 94.13 (11.39)</td><td></td></tr><tr><td>Ear length (E)</td><td>20-39; 31.81 (3.61)</td><td>27.65-31; 29.33 (2.37)</td></tr><tr><td>Weight (W)</td><td>16-55; 34.07 (7.48)</td><td></td></tr></tbody></table>

1253
1254 **Figure Legends**

1255
1256 **Figure 1.** Adult *Trachops cirrhosus* captured in Parque Nacional Yasuní, Orellana, Ecuador. Note
1257 the conspicuous finger-like dermal projections on the chin and lips. Photo: Rubén D. Jarrín.

1258
1259 **Figure 2.** Collecting localities of the *Trachops* specimens analyzed in this study. The localities are
1260 shown in the Supplementary Table 1. Black symbols represent *Trachops cirrhosus*, gray symbols
1261 represent *Trachops coffini*, and white symbols represent *Trachops ehrhardti*. Circles are
1262 specimens that were measured only, squares are sequenced only, and diamonds are both.

1263
1264 **Figure 3.** Maximum Likelihood phylogeny of *Trachops* based on complete mitochondrial genomes
1265 (nucleotide sequences). The tree was reconstructed in RAxML under the GTR+G+I model using 57
1266 complete mitogenomes including outgroups. Shaded semicircles on the nodes indicate ML
1267 bootstrap support (as a percentage) and Bayesian posterior probabilities from a congruent
1268 Bayesian Inference Analysis (see insert key).

1269
1270 **Figure 4.** Dorsal, ventral, and lateral views of the skulls and lateral view of the mandibles of
1271 *Trachops cirrhosus cirrhosus* (left); *T. c. coffini* (middle), and *T. c. ehrhardti* (right) Photos: Rubén
1272 D. Jarrín (QCAZ 11818); M. Alejandra Camacho (NMNH 244262); Anika Vogel (SMF 11718).

1273
1274 **Figure 5.** Principal components (PC's) from a PCA based on 13 measurements from 238
1275 individuals. Samples: *Trachops cirrhosus cirrhosus* (red circles), *T. c. coffini* (blue circles), and *T. c.
1276 *ehrhardti* (green circles).

1277
1278 **Figure 6.** Principal components analysis of 6 geographic groups and 238 specimens of *Trachops*
1279 of the 13 cranial measurements considered in the morphometric analysis. Meaning of Geographic
1280 abbreviation areas: **CAm** - Central America; **WSAm** - West South America; **ESAm** - East South
1281 America; NEC/NWV, NE Colombia and NW Venezuela; GS - the Guiana Shield; and AF - Atlantic Forest.

1282
1283
1284 **Figure 7.** Emended distributions of *Trachops cirrhosus* and *Trachops ehrhardti*. Black circles and crosses (• and ×) represent the recorded locations of *Trachops cirrhosus*. White circles and plus signs (o and+) correspond to the recorded locations of *Trachops ehrhardti*. The crosses and plus signs are the data collected by Fonseca (2019), sourced from https://github.com/bsf07/Defesa.git.

1287
1288
1289
1290 **FIGURES**

1291

1292 • These figures are only referential. All the figures are presented in TIFF format in separate.

1293

<figure><img src="image_1.png" /></figure>

1294

1295 **Figure 1.** Adult *Trachops cirrhosus* captured in Parque Nacional Yasuní, Orellana, Ecuador. Note

1296

the conspicuous finger-like dermal projections on the chin and lips. Photo: Rubén D. Jarrín.
<figure><img src="image_2.png" /><figcaption>**Figure 2.** Collecting localities of the *Trachops* specimens analyzed in this study. The localities are shown in the Supplementary Table 1. Black symbols represent *Trachops cirrhosus*, gray symbols represent *Trachops coffini*, and white symbols represent *Trachops ehrhardti*. Circles are specimens that were measured only, squares are sequenced only, and diamonds are both.</figcaption></figure>
<figure><img src="image_3.png" /><figcaption>**Figure 3.** Maximum Likelihood phylogeny of *Trachops* based on complete mitochondrial genomes (nucleotide sequences). The tree was reconstructed in RAxML under the GTR+G+I model using 57 complete mitogenomes including outgroups. Shaded semicircles on the nodes indicate ML bootstrap support (as a percentage) and Bayesian posterior probabilities from a congruent Bayesian Inference Analysis (see insert key).</figcaption></figure>
<figure><img src="image_4.png" /><figcaption>**Figure 4.** Dorsal, ventral, and lateral views of the skulls and lateral view of the mandibles of *Trachops cirrhosus cirrhosus* (left); *T. c. coffini* (middle), and *T. c. ehrhardti* (right) Photos: Rubén D. Jarrín (QCAZ 11818); Alejandra Camacho (NMNH 244262); Anika Vogel (SMF 11718).</figcaption></figure>

1310

1311

1312

1313

1314
<figure><img src="image_5.png" /></figure>

1315

1316

1317

1318

<caption>**Figure 5.** Principal components (PC's) from a PCA based on 13 measurements from 238 individuals. Samples: *Trachops cirrhosus cirrhosus* (red circles), *T. c. coffini* (blue circles), and *T. c. ehrhardti* (green circles).</caption>
<figure><img src="image_6.png" /><figcaption>**Figure 6.** Principal components analysis of 6 geographic groups and 238 specimens of *Trachops* of the 13 cranial measurements considered in the morphometric analysis. Meaning of Geographic abbreviation areas: **CAm** - Central America; **WSAm** - West South America; **ESAm** - East South America; **NEC/NWV**, NE Colombia and NW Venezuela; **GS** - the Guiana Shield; and **AF** - Atlantic Forest.</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>**Figure 7.** Emended distributions of *Trachops cirrhosus* and *Trachops ehrhardti*. Black circles and crosses (• and ×) represent the recorded locations of *Trachops cirrhosus*. White circles and plus signs (o and+) correspond to the recorded locations of *Trachops ehrhardti*. The crosses and plus signs are the data collected by Fonseca (2019), sourced from https://github.com/bsf07/Defesa.git.</figcaption></figure>

1325
1326
1327
1328
1329
1330
1331 **Appendix 1.** List of specimens used in the present study including voucher identification, 1332 GenBank accession number, taxonomic information, and general features of the mitogenome 1333 assemblies.

<table><thead><tr><th>Museum number</th><th>GenBank Accession number</th><th>Species</th><th>N° of reads</th><th>Mean coverage</th><th>N° of mtDNA reads</th><th>mtDNA length</th><th>GC content (%)</th></tr></thead><tbody><tr><td>AMNH-15103</td><td>Pending</td><td>T. cirrhosus</td><td>4324942</td><td>104,79</td><td>29909</td><td>16584</td><td>37,58</td></tr><tr><td>AMNH-31522</td><td>Pending</td><td>T. cirrhosus</td><td>2236998</td><td>37,01</td><td>10356</td><td>16601</td><td>37,58</td></tr><tr><td>AMNH-94538</td><td>Pending</td><td>T. cirrhosus</td><td>2151243</td><td>23,34</td><td>5747</td><td>16591</td><td>37,37</td></tr><tr><td>AMNH-129064</td><td>Pending</td><td>T. cirrhosus</td><td>2740862</td><td>79,92</td><td>22336</td><td>16595</td><td>37,58</td></tr><tr><td>AMNH-147767</td><td>Pending</td><td>T. cirrhosus</td><td>2789364</td><td>7,54</td><td>1891</td><td>16598</td><td>37,58</td></tr><tr><td>AMNH-185889</td><td>Pending</td><td>T. cirrhosus</td><td>3336996</td><td>17,44</td><td>4212</td><td>16596</td><td>37,25</td></tr><tr><td>AMNH-185891</td><td>Pending</td><td>T. cirrhosus</td><td>4357962</td><td>101,3</td><td>22996</td><td>16625</td><td>37,3</td></tr><tr><td>AMNH-230213</td><td>Pending</td><td>T. cirrhosus</td><td>4526938</td><td>399,14</td><td>90418</td><td>16622</td><td>37,66</td></tr><tr><td>AMNH-267935</td><td>Pending</td><td>T. cirrhosus</td><td>3494936</td><td>164,35</td><td>37044</td><td>16621</td><td>37,53</td></tr><tr><td>AMNH-272813</td><td>Pending</td><td>T. cirrhosus</td><td>5430224</td><td>147,99</td><td>33493</td><td>16655</td><td>37,58</td></tr><tr><td>AMNH-278476</td><td>Pending</td><td>T. cirrhosus</td><td>5304394</td><td>57,21</td><td>12932</td><td>16653</td><td>37,59</td></tr><tr><td>FMNH-72150</td><td>Pending</td><td>T. cirrhosus</td><td>7428142</td><td>7,64</td><td>1773</td><td>16578</td><td>37,35</td></tr><tr><td>FMNH-114877</td><td>Pending</td><td>T. cirrhosus</td><td>7529998</td><td>82,99</td><td>18741</td><td>16598</td><td>37,36</td></tr><tr><td>FMNH-174887</td><td>Pending</td><td>T. cirrhosus</td><td>5776246</td><td>1237,67</td><td>282126</td><td>16743</td><td>37,7</td></tr><tr><td>FMNH-174888</td><td>Pending</td><td>T. cirrhosus</td><td>5550234</td><td>282,11</td><td>64164</td><td>16713</td><td>37,68</td></tr><tr><td>MSB-311655</td><td>Pending</td><td>T. cirrhosus</td><td>4271430</td><td>68,59</td><td>15555</td><td>16600</td><td>37,58</td></tr><tr><td>MSB-311656</td><td>Pending</td><td>T. cirrhosus</td><td>4543408</td><td>261,75</td><td>59681</td><td>16602</td><td>37,58</td></tr><tr><td>MSB-311734</td><td>Pending</td><td>T. cirrhosus</td><td>4944460</td><td>169,89</td><td>38553</td><td>16603</td><td>37,37</td></tr><tr><td>MSB-311736</td><td>Pending</td><td>T. cirrhosus</td><td>5215900</td><td>62,19</td><td>14095</td><td>16603</td><td>37,37</td></tr><tr><td>QCAZ-2459</td><td>Pending</td><td>T. cirrhosus</td><td>3690346</td><td>54,62</td><td>12560</td><td>16630</td><td>37,46</td></tr><tr><td>QCAZ-4234</td><td>Pending</td><td>T. cirrhosus</td><td></td><td></td><td></td><td>16822</td><td>37,60</td></tr><tr><td>QCAZ-9239</td><td>Pending</td><td>T. cirrhosus</td><td>3751390</td><td>157,88</td><td>35638</td><td>16623</td><td>37,47</td></tr><tr><td>QCAZ-10617</td><td>Pending</td><td>T. cirrhosus</td><td>5240730</td><td>7,99</td><td>1829</td><td>16620</td><td>37,47</td></tr><tr><td>QCAZ-11632</td><td>Pending</td><td>T. cirrhosus</td><td>6075978</td><td>111,02</td><td>25064</td><td>16571</td><td>37,31</td></tr><tr><td>QCAZ-13317</td><td>Pending</td><td>T. cirrhosus</td><td></td><td></td><td></td><td>16619</td><td>37,50</td></tr><tr><td>QCAZ-13956</td><td>Pending</td><td>T. cirrhosus</td><td>5352738</td><td>29,08</td><td>6658</td><td>16621</td><td>37,35</td></tr><tr><td>QCAZ-14029</td><td>Pending</td><td>T. cirrhosus</td><td>6361566</td><td>104,18</td><td>23674</td><td>16619</td><td>37,42</td></tr><tr><td>QCAZ-17165</td><td>Pending</td><td>T. cirrhosus</td><td>6480538</td><td>20,02</td><td>4522</td><td>16614</td><td>37,34</td></tr><tr><td>QCAZ-18335</td><td>Pending</td><td>T. cirrhosus</td><td>4985186</td><td>83</td><td>18723</td><td>16579</td><td>37,35</td></tr><tr><td>ROM-95747</td><td>Pending</td><td>T. cirrhosus</td><td>5305082</td><td>299,46</td><td>67789</td><td>16624</td><td>37,32</td></tr><tr><td>ROM-98701</td><td>Pending</td><td>T. cirrhosus</td><td>5550346</td><td>43,24</td><td>9792</td><td>16612</td><td>37,67</td></tr><tr><td>ROM-99493</td><td>Pending</td><td>T. cirrhosus</td><td>3211086</td><td>79,6</td><td>18085</td><td>16621</td><td>37,31</td></tr><tr><td>ROM-103359</td><td>Pending</td><td>T. cirrhosus</td><td>4896612</td><td>14,83</td><td>3326</td><td>16317</td><td>36,24</td></tr><tr><td>ROM-103495</td><td>Pending</td><td>T. cirrhosus</td><td>4853726</td><td>522,78</td><td>118097</td><td>16621</td><td>37,59</td></tr><tr><td>ROM-104214</td><td>Pending</td><td>T. cirrhosus</td><td>4148188</td><td>39,36</td><td>9909</td><td>16582</td><td>37,3</td></tr><tr><td>ROM-107843</td><td>Pending</td><td>T. cirrhosus</td><td>4250814</td><td>17,87</td><td>4049</td><td>16617</td><td>37,41</td></tr></tbody></table>
<table><tr><td>ROM-107908</td><td>Pending</td><td>T. cirrhosus</td><td>3634508</td><td>67,38</td><td>15210</td><td>16593</td><td>37,4</td></tr><tr><td>ROM-107926</td><td>Pending</td><td>T. cirrhosus</td><td>3176256</td><td>169,41</td><td>38250</td><td>16602</td><td>37,41</td></tr><tr><td>ROM-107935</td><td>Pending</td><td>T. cirrhosus</td><td>3613554</td><td>260,21</td><td>58765</td><td>16607</td><td>37,39</td></tr><tr><td>ROM-107945</td><td>Pending</td><td>T. cirrhosus</td><td>4380050</td><td>281,34</td><td>63704</td><td>16637</td><td>37,65</td></tr><tr><td>ROM-115072</td><td>Pending</td><td>T. cirrhosus</td><td>6091173</td><td>10,53</td><td>2211</td><td>16577</td><td>37,29</td></tr><tr><td>ROM-115495</td><td>Pending</td><td>T. cirrhosus</td><td>7436128</td><td>21,57</td><td>4895</td><td>16624</td><td>37,42</td></tr><tr><td>ROM-117523</td><td>Pending</td><td>T. cirrhosus</td><td></td><td></td><td></td><td>16749</td><td>37,6</td></tr><tr><td>ROM-119312</td><td>Pending</td><td>T. cirrhosus</td><td>4225778</td><td>167,44</td><td>37808</td><td>16611</td><td>37,67</td></tr><tr><td>ROM-121044</td><td>Pending</td><td>T. cirrhosus</td><td>4673024</td><td>426,56</td><td>96388</td><td>16613</td><td>37,56</td></tr><tr><td>ROM-121090</td><td>Pending</td><td>T. cirrhosus</td><td>4051452</td><td>46,35</td><td>10501</td><td>16622</td><td>37,65</td></tr><tr><td>ROM-121159</td><td>Pending</td><td>T. cirrhosus</td><td>6174610</td><td>1006,61</td><td>227410</td><td>16594</td><td>37,62</td></tr><tr><td>ROM-121252</td><td>Pending</td><td>T. cirrhosus</td><td>4190554</td><td>197,92</td><td>44724</td><td>16615</td><td>37,69</td></tr><tr><td>ROM-122057</td><td>Pending</td><td>T. cirrhosus</td><td>4946730</td><td>215,59</td><td>49150</td><td>16622</td><td>37,67</td></tr><tr><td>ROM-125132</td><td>Pending</td><td>T. cirrhosus</td><td>5841756</td><td>217,79</td><td>48522</td><td>16631</td><td>37,65</td></tr><tr><td>ROM-125898</td><td>Pending</td><td>T. cirrhosus</td><td>5053904</td><td>26,77</td><td>6039</td><td>16589</td><td>37,51</td></tr><tr><td>ROM-125914</td><td>Pending</td><td>T. cirrhosus</td><td>5364036</td><td>132,02</td><td>29962</td><td>16606</td><td>37,53</td></tr><tr><td>FMNH-94717</td><td>Pending</td><td>T. erhardti</td><td>3597194</td><td>8,72</td><td>1996</td><td>16618</td><td>37,03</td></tr><tr><td>FMNH-141595</td><td>Pending</td><td>T. erhardti</td><td>4767214</td><td>20,62</td><td>4716</td><td>16619</td><td>37,3</td></tr><tr><td></td><td></td><td>Average</td><td>4731900,47</td><td>138,74</td><td>36649,90</td><td>16615,22</td><td>37,46</td></tr><tr><td></td><td></td><td>Standard deviation</td><td>1242366,47</td><td>172,72</td><td>52650,26</td><td>60,37</td><td>0,22</td></tr></table>
<figure><img src="image_8.png" /><figcaption>Adult Trachops cirrhosus captured in Parque Nacional Yasuní, Orellana, Ecuador. Note the conspicuous finger-like dermal projections on the chin and lips. Photo: Rubén D. Jarrín.</figcaption></figure>

649x979mm (28 x 28 DPI)
<figure><img src="image_9.png" /></figure>

Collecting localities of the Trachops specimens analyzed in this study. The localities are shown in the Supplementary Table 1. Black symbols represent Trachops cirrhosus, gray symbols represent Trachops coffini, and white symbols represent Trachops ehrhardti. Circles are specimens that were measured only, squares are sequenced only, and diamonds are both.

1407x979mm (28 x 28 DPI)
<figure><img src="image_10.png" /></figure>

Maximum Likelihood phylogeny of Trachops based on complete mitochondrial genomes (nucleotide sequences). The tree was reconstructed in RAxML under the GTR+G+I model using 57 complete mitogenomes including outgroups. Shaded semicircles on the nodes indicate ML bootstrap support (as a percentage) and Bayesian posterior probabilities from a congruent Bayesian Inference Analysis (see insert key).

1041x910mm (28 x 28 DPI)
<figure><img src="image_11.png" /><figcaption>Dorsal, ventral, and lateral views of the skulls and lateral view of the mandibles of Trachops cirrhosus cirrhosus (left); T. c. coffini (middle), and T. c. ehrhardti (right) Photos: Rubén D. Jarrín (QCAZ 11818); M. Alejandra Camacho (NMNH 244262); Anika Vogel (SMF 11718).</figcaption></figure>

1415x979mm (28 x 28 DPI)
<figure><img src="image_12.png" /></figure>

Principal components (PC's) from a PCA based on 13 measurements from 238 individuals. Samples: *Trachops cirrhosus cirrhosus* (red circles), *T. c. coffini* (blue circles), and *T. c. ehrhardti* (green circles).

1469x979mm (28 x 28 DPI)
<figure><img src="image_13.png" /></figure>

Principal components analysis of 6 geographic groups and 238 specimens of Trachops of the 13 cranial measurements considered in the morphometric analysis. Meaning of Geographic abbreviation areas: CAm - Central America; WSAm - West South America; ESAm - East South America; NEC/NWV, NE Colombia and NW Venezuela; GS - the Guiana Shield; and AF - Atlantic Forest.

1469x979mm (28 x 28 DPI)
<figure><img src="image_14.png" /></figure>

Emended distributions of Trachops cirrhosus and Trachops ehrhardti. Black circles and crosses (• and □) represent the recorded locations of Trachops cirrhosus. White circles and plus signs (□ and □) correspond to the recorded locations of Trachops ehrhardti. The crosses and plus signs are the data collected by Fonseca (2019), sourced from https://github.com/bsf07/Defesa.git.

199x139mm (300 x 300 DPI)
<table><caption>Supplementary Table 1. Specimens, samples, biorepositories and geographic information.</caption><thead><tr><th>Museum</th><th># voucher</th><th>Unique</th><th>Measured</th><th>Treatment</th><th>Species</th></tr></thead><tbody><tr><td>ROM</td><td>120475</td><td>ROM-120475</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>278284</td><td>AMNH-278284</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>278402</td><td>AMNH-278402</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>278668</td><td>AMNH-278668</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>278748</td><td>AMNH-278748</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>583009</td><td>USNM-583009</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>209349</td><td>AMNH-209349</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>209350</td><td>AMNH-209350</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>209348</td><td>AMNH-209348</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>209351</td><td>AMNH-209351</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>210679</td><td>AMNH-210679</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>210680</td><td>AMNH-210680</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>210681</td><td>AMNH-210681</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>114877</td><td>FMNH-114877</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>584479</td><td>USNM-584479</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>530956</td><td>USNM-530956</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>530957</td><td>USNM-530957</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>92252</td><td>AMNH-92252</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>94536</td><td>AMNH-94536</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>94537</td><td>AMNH-94537</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>549355</td><td>USNM-549355</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>549354</td><td>USNM-549354</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>361534</td><td>USNM-361534</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>361533</td><td>USNM-361533</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>460090</td><td>USNM-460090</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>94538</td><td>AMNH-94538</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>95500</td><td>AMNH-95500</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>235204</td><td>FMNH-235204</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>SMF</td><td>11716</td><td>SMF-11716</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>SMF</td><td>11717</td><td>SMF-11717</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>SMF</td><td>11718</td><td>SMF-11718</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>200992</td><td>USNM-200992</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>256301</td><td>AMNH-256301</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>141595</td><td>FMNH-141595</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>94717</td><td>FMNH-94717</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>954</td><td>IAvH-M-954</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>1331</td><td>IAvH-M-1331</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>499295</td><td>USNM-499295</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>499764</td><td>USNM-499764</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>9466</td><td>IAvH-M-9466</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>9628</td><td>IAvH-M-9628</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr></tbody></table>
<table><tr><td>IAvH-M</td><td>9470</td><td>IAvH-M-9470</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>10098</td><td>IAvH-M-10098</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>9976</td><td>IAvH-M-9976</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>92309</td><td>FMNH-92309</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>92311</td><td>FMNH-92311</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281054</td><td>USNM-281054</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>7141</td><td>IAvH-M-7141</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>10622</td><td>IAvH-M-10622</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>9970</td><td>IAvH-M-9970</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>7850</td><td>IAvH-M-7850</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>8109</td><td>IAvH-M-8109</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>8110</td><td>IAvH-M-8110</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281926</td><td>USNM-281926</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281927</td><td>USNM-281927</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281049</td><td>USNM-281049</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281050</td><td>USNM-281050</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281037</td><td>USNM-281037</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281038</td><td>USNM-281038</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281039</td><td>USNM-281039</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281040</td><td>USNM-281040</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>281045</td><td>USNM-281045</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>11372</td><td>IAvH-M-11372</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>UV</td><td>2745</td><td>UV-2745</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>UV</td><td>2746</td><td>UV-2746</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>15103</td><td>AMNH-15103</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>2218</td><td>IAvH-M-2218</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>11316</td><td>IAvH-M-11316</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>88070</td><td>FMNH-88070</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>UV</td><td>5549</td><td>UV-5549</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>142158</td><td>AMNH-142158</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>UV</td><td>6996</td><td>UV-6996</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>255713</td><td>AMNH-255713</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>71533</td><td>FMNH-71533</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>72150</td><td>FMNH-72150</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>UV</td><td>5550</td><td>UV-5550</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>2921</td><td>IAvH-M-2921</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>8332</td><td>IAvH-M-8332</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>IAvH-M</td><td>8333</td><td>IAvH-M-8333</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>562764</td><td>USNM-562764</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>562763</td><td>USNM-562763</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>565808</td><td>USNM-565808</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>568489</td><td>USNM-568489</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>137225</td><td>FMNH-137225</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr></table>
<table><tr><td>USNM</td><td>526241</td><td>USNM-526241</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>124102</td><td>FMNH-124102</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>9239</td><td>QCAZ-9239</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>5676</td><td>MEPN-5676</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>9237</td><td>QCAZ-9237</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>2341</td><td>QCAZ-2341</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>9238</td><td>QCAZ-9238</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>2014</td><td>MEPN-2014</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>3014</td><td>MEPN-3014</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>18335</td><td>QCAZ-18335</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>528489</td><td>USNM-528489</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>17164</td><td>QCAZ-17164</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>17165</td><td>QCAZ-17165</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>285</td><td>QCAZ-285</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>3290</td><td>QCAZ-3290</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>11818</td><td>QCAZ-11818</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>5603</td><td>MEPN-5603</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>18016</td><td>QCAZ-18016</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13876</td><td>QCAZ-13876</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13877</td><td>QCAZ-13877</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13415</td><td>QCAZ-13415</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13596</td><td>QCAZ-13596</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>7937</td><td>QCAZ-7937</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>8171</td><td>QCAZ-8171</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>8172</td><td>QCAZ-8172</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>8079</td><td>QCAZ-8079</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13956</td><td>QCAZ-13956</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>14029</td><td>QCAZ-14029</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>3289</td><td>QCAZ-3289</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>1116</td><td>QCAZ-1116</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>5125</td><td>MEPN-5125</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>4477</td><td>MEPN-4477</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>1475</td><td>QCAZ-1475</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>10617</td><td>QCAZ-10617</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>4234</td><td>QCAZ-4234</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>2013</td><td>MEPN-2013</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>4555</td><td>MEPN-4555</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MEPN</td><td>4482</td><td>MEPN-4482</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>7228</td><td>QCAZ-7228</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>6850</td><td>QCAZ-6850</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>6855</td><td>QCAZ-6855</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>6870</td><td>QCAZ-6870</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>13317</td><td>QCAZ-13317</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr></table>
<table><tr><td>QCAZ</td><td>11632</td><td>QCAZ-11632</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>7051</td><td>QCAZ-7051</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>7578</td><td>QCAZ-7578</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>10172</td><td>QCAZ-10172</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>587</td><td>QCAZ-587</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>5070</td><td>QCAZ-5070</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>QCAZ</td><td>2459</td><td>QCAZ-2459</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267935</td><td>AMNH-267935</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>266079</td><td>AMNH-266079</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267926</td><td>AMNH-267926</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267928</td><td>AMNH-267928</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267930</td><td>AMNH-267930</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267934</td><td>AMNH-267934</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>267936</td><td>AMNH-267936</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>266080</td><td>AMNH-266080</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>266089</td><td>AMNH-266089</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>65184</td><td>FMNH-65184</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>99493</td><td>ROM-99493</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244252</td><td>USNM-244252</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244254</td><td>USNM-244254</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244256</td><td>USNM-244256</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244262</td><td>USNM-244262</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244264</td><td>USNM-244264</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244266</td><td>USNM-244266</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244268</td><td>USNM-244268</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>244269</td><td>USNM-244269</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>582290</td><td>USNM-582290</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>98701</td><td>ROM-98701</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>115072</td><td>ROM-115072</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>115495</td><td>ROM-115495</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>122057</td><td>ROM-122057</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>125132</td><td>ROM-125132</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>582292</td><td>USNM-582292</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>582291</td><td>USNM-582291</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>103359</td><td>ROM-103359</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>103495</td><td>ROM-103495</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>182717</td><td>AMNH-182717</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>119312</td><td>ROM-119312</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>337270</td><td>USNM-337270</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>339719</td><td>USNM-339719</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>339720</td><td>USNM-339720</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>337289</td><td>USNM-337289</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129065</td><td>AMNH-129065</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr></table>
<table><tr><td>AMNH</td><td>129072</td><td>AMNH-129072</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129080</td><td>AMNH-129080</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129081</td><td>AMNH-129081</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129090</td><td>AMNH-129090</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129732</td><td>AMNH-129732</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129736</td><td>AMNH-129736</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129064</td><td>AMNH-129064</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>129730</td><td>AMNH-129730</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>95747</td><td>ROM-95747</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185891</td><td>AMNH-185891</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185886</td><td>AMNH-185886</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185889</td><td>AMNH-185889</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185887</td><td>AMNH-185887</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185888</td><td>AMNH-185888</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>185890</td><td>AMNH-185890</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>332697</td><td>USNM-332697</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>332698</td><td>USNM-332698</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>332699</td><td>USNM-332699</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>332700</td><td>USNM-332700</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>554264</td><td>USNM-554264</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>464596</td><td>USNM-464596</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>464597</td><td>USNM-464597</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>104214</td><td>ROM-104214</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>311960</td><td>USNM-311960</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>MSB</td><td>311736</td><td>MSB-311736</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>MSB</td><td>311734</td><td>MSB-311734</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>MSB</td><td>311655</td><td>MSB-311655</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>MSB</td><td>311656</td><td>MSB-311656</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309362</td><td>USNM-309362</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309364</td><td>USNM-309364</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309367</td><td>USNM-309367</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309368</td><td>USNM-309368</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309369</td><td>USNM-309369</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309370</td><td>USNM-309370</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>178902</td><td>AMNH-178902</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>143757</td><td>AMNH-143757</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>147767</td><td>AMNH-147767</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>42564</td><td>AMNH-42564</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>575470</td><td>USNM-575470</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>312982</td><td>USNM-312982</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>312983</td><td>USNM-312983</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>312984</td><td>USNM-312984</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>312985</td><td>USNM-312985</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr></table>
<table><tr><td>USNM</td><td>174884</td><td>USNM-174884</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>305199</td><td>USNM-305199</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>305200</td><td>USNM-305200</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>305201</td><td>USNM-305201</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309178</td><td>USNM-309178</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309179</td><td>USNM-309179</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309180</td><td>USNM-309180</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>309181</td><td>USNM-309181</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>92672</td><td>FMNH-92672</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>233227</td><td>AMNH-233227</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>236002</td><td>AMNH-236002</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>272813</td><td>AMNH-272813</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>273176</td><td>AMNH-273176</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>272820</td><td>AMNH-272820</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>272821</td><td>AMNH-272821</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>74026</td><td>AMNH-74026</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>278476</td><td>AMNH-278476</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>76162</td><td>AMNH-76162</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>76163</td><td>AMNH-76163</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>76164</td><td>AMNH-76164</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>13277</td><td>AMNH/MUSM-13277</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>13278</td><td>AMNH/MUSM-13278</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>13279</td><td>AMNH/MUSM-13279</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>13280</td><td>AMNH/MUSM-13280</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>15286</td><td>AMNH/MUSM-15286</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH/MUSM</td><td>15287</td><td>AMNH/MUSM-15287</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>566489</td><td>USNM-566489</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>566490</td><td>USNM-566490</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>566491</td><td>USNM-566491</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>174887</td><td>FMNH-174887</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>FMNH</td><td>174888</td><td>FMNH-174888</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>230213</td><td>AMNH-230213</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>364272</td><td>USNM-364272</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>125898</td><td>ROM-125898</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>125914</td><td>ROM-125914</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>126156</td><td>ROM-126156</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>117523</td><td>ROM-117523</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>121252</td><td>ROM-121252</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>121044</td><td>ROM-121044</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>121090</td><td>ROM-121090</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>121159</td><td>ROM-121159</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>175602</td><td>AMNH-175602</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>256313</td><td>AMNH-256313</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>180034</td><td>AMNH-180034</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr></table>
<table><tr><td>AMNH</td><td>175603</td><td>AMNH-175603</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>175600</td><td>AMNH-175600</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>107843</td><td>ROM-107843</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>388865</td><td>USNM-388865</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>388869</td><td>USNM-388869</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>407611</td><td>USNM-407611</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>407614</td><td>USNM-407614</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>373534</td><td>USNM-373534</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>373539</td><td>USNM-373539</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>235556</td><td>AMNH-235556</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>235557</td><td>AMNH-235557</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>235551</td><td>AMNH-235551</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>517312</td><td>USNM-517312</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>107935</td><td>ROM-107935</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>107908</td><td>ROM-107908</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>107926</td><td>ROM-107926</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>388852</td><td>USNM-388852</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>388856</td><td>USNM-388856</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>444405</td><td>USNM-444405</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>385510</td><td>USNM-385510</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>385511</td><td>USNM-385511</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>31522</td><td>AMNH-31522</td><td>Yes</td><td>Both</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>31520</td><td>AMNH-31520</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>AMNH</td><td>31521</td><td>AMNH-31521</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>444404</td><td>USNM-444404</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>371556</td><td>USNM-371556</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>371557</td><td>USNM-371557</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>419151</td><td>USNM-419151</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>419153</td><td>USNM-419153</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>419242</td><td>USNM-419242</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>444396</td><td>USNM-444396</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>444398</td><td>USNM-444398</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>ROM</td><td>107945</td><td>ROM-107945</td><td>No</td><td>Sequenced</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>444382</td><td>USNM-444382</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>371559</td><td>USNM-371559</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>371560</td><td>USNM-371560</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr><tr><td>USNM</td><td>419248</td><td>USNM-419248</td><td>Yes</td><td>Measured</td><td>Trachops cirrhosus</td></tr></table>

TOTALS

238

22

Both

216

Measured

56

Sequenced
<table><thead><tr><td>subespecies</td><td>Country</td><td>State/Province</td></tr></thead><tbody><tr><td>coffini</td><td>Belize</td><td>Orange Walk County</td></tr><tr><td>coffini</td><td>Belize</td><td>Orange Walk District</td></tr><tr><td>coffini</td><td>Belize</td><td>Orange Walk District</td></tr><tr><td>coffini</td><td>Belize</td><td>Orange Walk District</td></tr><tr><td>coffini</td><td>Belize</td><td>Orange Walk District</td></tr><tr><td>coffini</td><td>Belize</td><td>Stann Creek</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Beni</td></tr><tr><td>cirrhosus</td><td>Bolivia</td><td>Santa Cruz</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Amazonas</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Amazonas</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Amazonas</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Brazil</td><td>Rondonia</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Santa Catarina</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Santa Catarina</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Santa Catarina</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Santa Catarina</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Sao Paulo</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Sao Paulo</td></tr><tr><td>ehrhardti</td><td>Brazil</td><td>Sao Paulo</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Antioquia</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Antioquia</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Antioquia</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Antioquia</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr></tbody></table>
<table><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Arauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Bolivar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Caquetá</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casanare</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casanare</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casanare</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casanare</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Casar</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Córdoba</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Guainía</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Guainía</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Magdalena</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Meta</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Meta</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Meta</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Meta</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Meta</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Nariño</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Norte de Santander</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Putumayo</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Putumayo</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Valle del Cauca</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Vichada</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Vichada</td></tr><tr><td>cirrhosus</td><td>Colombia</td><td>Vichada</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Guanacaste</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Heredia</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Heredia</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Limon</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Limón</td></tr></table>
<table><tr><td>cirrhosus</td><td>Costa Rica</td><td>Puntarenas</td></tr><tr><td>cirrhosus</td><td>Costa Rica</td><td>Puntarenas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Esmeraldas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Guayas</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Los Ríos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Los Ríos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Los Ríos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Napo</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Napo</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Napo</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Napo</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Orellana</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Pastaza</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Pichincha</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbios</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbios</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbios</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr></table>
<table><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Sucumbíos</td></tr><tr><td>cirrhosus</td><td>Ecuador</td><td>Zamora Chinchipe</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Cayenne</td></tr><tr><td>cirrhosus</td><td>French Guiana</td><td>Paracou</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Chimaltenango</td></tr><tr><td>coffini</td><td>Guatemala</td><td>El Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>coffini</td><td>Guatemala</td><td>Peten</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Barima-Waini</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Barima-Waini</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Cuyuni-Mazaruni</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Essequibo Islands-West</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Potaro-Siparuni</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Potaro-Siparuni</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Demerara-Berbice</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Demerara-Berbice</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Demerara-Berbice</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Demerara-Berbice</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>cirrhosus</td><td>Guyana</td><td>Upper Takutu - Upper Es</td></tr><tr><td>coffini</td><td>Honduras</td><td>Lempira</td></tr></table>
<table><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Panama</td><td></td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Huanuco</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Huanuco</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Loreto</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Madre De Dios</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Madre De Dios</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Madre De Dios</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Madre de Dios</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Madre de Dios</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Pasco</td></tr><tr><td>cirrhosus</td><td>Peru</td><td>Pasco</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Brokopondo</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Brokopondo</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Para</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Sipaliwini</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Sipaliwini</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Sipaliwini</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Sipaliwini</td></tr><tr><td>cirrhosus</td><td>Suriname</td><td>Sipaliwini</td></tr><tr><td>cirrhosus</td><td>Trinidad and T</td><td>Trinidad</td></tr><tr><td>cirrhosus</td><td>Trinidad and T</td><td>Trinidad</td></tr><tr><td>cirrhosus</td><td>Trinidad and T</td><td>Trinidad</td></tr></table>
<table><thead><tr><td>Verbatim Precise Locality</td><td>Latitude</td><td>Longitude</td></tr></thead><tbody><tr><td>Lamanai Lodge</td><td>17.752072</td><td>-88.656762</td></tr><tr><td>Ka'Kabish Archaeological Reserve</td><td>17.767250</td><td>-88.867436</td></tr><tr><td>Ka'Kabish Archaeological Reserve</td><td>17.767250</td><td>-88.867436</td></tr><tr><td>Ka'Kabish Archaeological Reserve</td><td>17.767250</td><td>-88.867436</td></tr><tr><td>Ka'Kabish Archaeological Reserve</td><td>17.767250</td><td>-88.867436</td></tr><tr><td>Cockscomb Basin Wildlife Sanctuary</td><td>16.773611</td><td>-88.532500</td></tr><tr><td>ca. 1.5 kilometers below Costa Marquez, Itenez River</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>ca. 1.5 kilometers below Costa Marquez, Itenez River</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>Baures River mouth</td><td>-11.094083</td><td>-65.281711</td></tr><tr><td>ca. 4 kilometers above Costa Marques, Itenez River</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>ca. 5 kilometers northwest of Grande River mouth</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>ca. 5 kilometers northwest of Grande River mouth</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>ca. 5 kilometers northwest of Grande River mouth</td><td>-13.334992</td><td>-63.439769</td></tr><tr><td>El Beni, Mamore, Camino Vilches</td><td>-13.066700</td><td>-64.800000</td></tr><tr><td>National Park Kempff Mercado, El Refugio</td><td>-14.766389</td><td>-61.034722</td></tr><tr><td>Manaus, 80 Km N (By Road)</td><td>-2.510833</td><td>-60.017500</td></tr><tr><td>Manaus, 80 Km N (By Road)</td><td>-2.510833</td><td>-60.017500</td></tr><tr><td>Rosarinho. Rio Madeira</td><td>-5.8753</td><td>-61.3903</td></tr><tr><td>Tauari, Tapajos River</td><td>-3.083333</td><td>-55.100000</td></tr><tr><td>Tauari, Tapajos River</td><td>-3.083333</td><td>-55.100000</td></tr><tr><td>Altamira, 17 Km S (By Road), Caverna Do Tatajuba</td><td>-3.630000</td><td>-52.350000</td></tr><tr><td>Altamira, 52 Km SSW, E Bank Rio Xingu</td><td>-3.650000</td><td>-52.370000</td></tr><tr><td>Belem, Fazenda Velha</td><td>-1.444722</td><td>-48.501389</td></tr><tr><td>Belem, Station A Ian</td><td>-1.444722</td><td>-48.501389</td></tr><tr><td>Belem, Varzea</td><td>-1.444722</td><td>-48.501389</td></tr><tr><td>Tapajos River, Igarape Brabo</td><td>-2.466667</td><td>-54.750000</td></tr><tr><td>Tapajos River, Inajatuba</td><td>-4.115556</td><td>-56.609722</td></tr><tr><td>Rondonia, Cachoeira Nazare, Rio Ji-Parana</td><td>-8.225000</td><td>-62.726667</td></tr><tr><td>Joinville</td><td>-26.293056</td><td>-48.827778</td></tr><tr><td>Joinville</td><td>-26.293056</td><td>-48.827778</td></tr><tr><td>Joinville</td><td>-26.293056</td><td>-48.827778</td></tr><tr><td>Mount Hansa</td><td>-26.881944</td><td>-48.451389</td></tr><tr><td>Bairro do Betari, Gruta do Betari</td><td>-24.586389</td><td>-48.593889</td></tr><tr><td>Ilha do Cardoso.</td><td>-25.133300</td><td>-47.966700</td></tr><tr><td>Primeiro Morro.</td><td>-24.366700</td><td>-47.816700</td></tr><tr><td>25 Km South, 22 Km West from Zaragoza, La Tirana</td><td>7.261694</td><td>-75.028628</td></tr><tr><td>26 Km South, 22 Km West from Zaragoza, Aljibes</td><td>7.258047</td><td>-75.047181</td></tr><tr><td>Zaragoza, 25 Km S, 22 Km W, At La Tirana</td><td>7.485089</td><td>-74.867306</td></tr><tr><td>Zaragoza, 25 Km S, 22 Km W, At La Tirana</td><td>7.485089</td><td>-74.867306</td></tr><tr><td>Finca La Tormenta, Caño Salas</td><td>6.812111</td><td>-71.0755</td></tr><tr><td>Caño Juriepe, Finca La Aurora, Bosques de Juriepe</td><td>6.2339722</td><td>-69.716333</td></tr></tbody></table>
<table><tr><td>Reserva Forestal Protectora Rio Tame, Vereda Sabana de La V</td><td>6.3843611</td><td>-71.926389</td></tr><tr><td>Vereda Puna Puna, Hacienda la Envidia</td><td>6.3301667</td><td>-71.772222</td></tr><tr><td>Vereda San Joaquín, finca El Santuario, bosque de vega, cerca</td><td>6.222532</td><td>-71.566834</td></tr><tr><td>Arauca, 1000ft: Rio Arauca.</td><td>6.203630</td><td>-68.964500</td></tr><tr><td>Arauca, 1000ft: Rio Arauca.</td><td>6.203630</td><td>-68.964500</td></tr><tr><td>Mompos, Norosi</td><td>9.239653</td><td>-74.425617</td></tr><tr><td>PNN Serrania de Chiribiquete, Corregimiento SE río Mesay, P</td><td>0.0741667</td><td>-72.451389</td></tr><tr><td>Vereda Cupiagua, caño abajo de pozo V</td><td>5.2262222</td><td>-72.626858</td></tr><tr><td>Vereda La Manga, finca La Reserva, caño Yaguarapo</td><td>6.198954</td><td>-71.603834</td></tr><tr><td>Corregimiento La Hermosa, finca Nicaragua, bajo de quebrad</td><td>5.6641667</td><td>-70.2425</td></tr><tr><td>Sin descripción del lugar de colecta-Paz de Ariporo</td><td>5</td><td>-70</td></tr><tr><td>Sin descripción del lugar de colecta-Paz de Ariporo</td><td>5</td><td>-70</td></tr><tr><td>Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Valledupar, El Orinoco, Rio Cesar</td><td>9.474669</td><td>-74.058406</td></tr><tr><td>Valledupar, El Orinoco, Rio Cesar</td><td>9.474669</td><td>-74.058406</td></tr><tr><td>Valledupar, Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Valledupar, Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Valledupar, Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Valledupar, Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Valledupar, Rio Guaimaral</td><td>9.204211</td><td>-73.656131</td></tr><tr><td>Corregimiento El Porveni, Vereda El Peñon, Predio de ODL</td><td>9.3963611000</td><td>-75.7071944000</td></tr><tr><td>Randal Alto (ca. Zancudo). Río Inírida, Guainía</td><td>2.766667</td><td>-69.366667</td></tr><tr><td>Randal Alto (ca. Zancudo). Río Inírida, Guainía</td><td>2.766667</td><td>-69.366667</td></tr><tr><td>Bonda</td><td>11.233425</td><td>-74.123428</td></tr><tr><td>Trocha 1 Km de la Cabaña Duda</td><td>2.180033</td><td>-73.783108</td></tr><tr><td>Rivera del río Tillavá</td><td>3.8275556000</td><td>-71.2658417000</td></tr><tr><td>Meta, Villavicencio, 400m, San Juan de Arama, Los Micos.</td><td>3.283330</td><td>-73.883300</td></tr><tr><td>Quebrada Alegría, Fuente de Materiales Valle</td><td>3.717222</td><td>-74.296667</td></tr><tr><td>Villavicencio, 650m.</td><td>4.141944</td><td>-73.626667</td></tr><tr><td>Inguapí del Guadual, 15 km Tumaco</td><td>1.700000</td><td>-78.766667</td></tr><tr><td>Rio de Oro</td><td>8.294119</td><td>-73.387344</td></tr><tr><td>Putumayo, San Antonio, 185m: Rio Mecaya.</td><td>0.466667</td><td>-75.333300</td></tr><tr><td>Putumayo, San Antonio, 185m: Rio Mecaya.</td><td>0.466667</td><td>-75.333300</td></tr><tr><td>Fuente de materiales, Bahía de Málaga, 40 m Valle</td><td>3.976111</td><td>-77.325278</td></tr><tr><td>PNN El Tuparro, Centro Administrativo</td><td>5.309489</td><td>-67.894906</td></tr><tr><td>PNN El Tuparro, Corregimiento Santa Rita</td><td>5.286408</td><td>-68.319869</td></tr><tr><td>PNN El Tuparro, Corregimiento Santa Rita</td><td>5.286408</td><td>-68.319869</td></tr><tr><td>Refugio Rafael Rodriguez, Hacienda Palo Verde</td><td>10.350056</td><td>-85.349583</td></tr><tr><td>Parque Nacional Braulio Carrillo, San Miguel, 1 Km S, 11.5 Km</td><td>9.980775</td><td>-84.051450</td></tr><tr><td>River Station, Bridge At, La Selva</td><td>10.427456</td><td>-84.006339</td></tr><tr><td>Cano Palma Biological Station</td><td>10.600000</td><td>-83.530000</td></tr><tr><td>Limon, 10ft: Tortuguero, shore opposite.</td><td>10.583300</td><td>-83.516700</td></tr></table>
<table><tr><td>Osa Peninsula, Llorona, Piedra El Arco</td><td>9.401969</td><td>-84.080308</td></tr><tr><td>Puntarenas, Monteverde.</td><td>10.300000</td><td>-84.700000</td></tr><tr><td>Comuna San Francisco de Bogota</td><td>1.09343</td><td>-78.7063</td></tr><tr><td>Estero El Salto, Centro Comunal Awa Mataje</td><td>1.200000</td><td>-78.561667</td></tr><tr><td>San Lorenzo</td><td>1.275994</td><td>-78.832183</td></tr><tr><td>San Lorenzo, Estación La Chiquita</td><td>1.23333</td><td>-78.76</td></tr><tr><td>San Lorenzo, Estación La Chiquita</td><td>1.23333</td><td>-78.76</td></tr><tr><td>Urbina</td><td>1.036667</td><td>-78.773611</td></tr><tr><td>3 km al S de San Miguel (selva)</td><td></td><td></td></tr><tr><td>Central Hidroeléctirca Marcel Lianado de Wind-Carlas Julio-R</td><td>-0.9252312</td><td>-79.761266</td></tr><tr><td>Santo Domingo, 47 Km S (By Road), Rio Palenque Science Cer</td><td>-0.586400</td><td>-79.364700</td></tr><tr><td>Centro Científico Río Palenque</td><td>-0.58884</td><td>-79.36106</td></tr><tr><td>Centro Científico Río Palenque</td><td>-0.58884</td><td>-79.36106</td></tr><tr><td>Jondachi</td><td>-0.79215</td><td>-77.7772</td></tr><tr><td>Jumandi, 4 km N de Archidona</td><td>-0.8721</td><td>-77.7914</td></tr><tr><td>Gareno, Río Gareno, junto pozo de exploracion Nemora, terra</td><td>-1.0485</td><td>-77.3774</td></tr><tr><td>5 km SW de Marian</td><td>-0.083333</td><td>-76.350000</td></tr><tr><td>Amo C, 101,1 km S de Pompeya Sur</td><td>-0.9419245</td><td>-76.22474</td></tr><tr><td>Parque Nacional Yasuní, Tambococha</td><td>-1.02253</td><td>-75.448</td></tr><tr><td>Parque Nacional Yasuní, Tambococha</td><td>-1.02253</td><td>-75.448</td></tr><tr><td>Parroquia Cononaco, Bloque 16, Repsol YPF-NPF, Km 66, La C</td><td>-0.806932</td><td>-76.3976</td></tr><tr><td>Parroquia Cononaco, Territorio Huaorani, Zona Intangible, fr</td><td>-1.2671</td><td>-76.3836</td></tr><tr><td>Puente de Dosel Chuncho-Yura</td><td>-0.643669</td><td>-75.9102</td></tr><tr><td>Puente de Dosel Pandu</td><td>-0.684732</td><td>-75.9236</td></tr><tr><td>Puente de Dosel Pandu</td><td>-0.684732</td><td>-75.9236</td></tr><tr><td>Vía Nenke-Apaika, orillas del Río Huiririma</td><td>-0.64833</td><td>-75.9136</td></tr><tr><td>Estación Científica Yasuní, Sendero Barizo</td><td>-0.67505</td><td>-76.3999</td></tr><tr><td>Estación Científica Yasuní, Sendero Ceiba, junto a la estación</td><td>-0.67616</td><td>-76.3959</td></tr><tr><td>Parque Nacional Yasuní, Estación Científica Yasuní</td><td>-0.678193</td><td>-76.3966</td></tr><tr><td>Yasuní, P.N., El Saladero, km 78</td><td>-0.8333333</td><td>-76.333333</td></tr><tr><td>Caimitoyacu. Cerca de la boca de Suno</td><td></td><td></td></tr><tr><td>Iripari</td><td></td><td></td></tr><tr><td>El Edén, Río Napo bajo, cerca a la bocana del Río Yuturi</td><td>-0.539787</td><td>-76.0237</td></tr><tr><td>Comunidad Tarangaro</td><td>-1.40062</td><td>-77.3835</td></tr><tr><td>La Unión del Toachi</td><td>-0.31383</td><td>-78.9544</td></tr><tr><td>Reserva Biológica Limoncocha</td><td>-0.400000</td><td>-76.633333</td></tr><tr><td>Pozo petrolero Paujil</td><td></td><td></td></tr><tr><td>Río Aguarico. Comunidad Cofán. Zábalo</td><td>-0.400000</td><td>-75.633333</td></tr><tr><td>Destacamento Patria, hito 63</td><td>-0.46503</td><td>-75.3451</td></tr><tr><td>Puente del Río Cuyabeno, Hostería Monte Tour, Vía Tarapoa</td><td>-0.0315</td><td>-76.32111</td></tr><tr><td>Puente del Río Cuyabeno, Hostería Monte Tour, Vía Tarapoa</td><td>-0.0315</td><td>-76.32111</td></tr><tr><td>Puente del Río Cuyabeno, Hostería Monte Tour, Vía Tarapoa</td><td>-0.0315</td><td>-76.32111</td></tr><tr><td>Reserva de Producción Faunística Cuyabeno, Santa Elena</td><td>-0.315235</td><td>-75.9624</td></tr></table>
<table><tr><td>Reserva de Producción Faunística Cuyabeno, Jamu Lodge, ori</td><td>-0.034808</td><td>-76.1644</td></tr><tr><td>Zábalo</td><td>-0.31812</td><td>-75.7662</td></tr><tr><td>Zábalo</td><td>-0.31812</td><td>-75.7662</td></tr><tr><td>Nueva Esmeralda</td><td>-0.18383</td><td>-76.7334</td></tr><tr><td>Reserva Biológica Limoncocha</td><td>-0.4079</td><td>-76.6206</td></tr><tr><td>Shushufindi</td><td>-0.18329</td><td>-76.6451</td></tr><tr><td>Destacamento Shaime, Ríos Shaime y Nangaritza</td><td>-4.3344</td><td>-78.6807</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Paracou near Sinnamary</td><td>5.283333</td><td>-52.916667</td></tr><tr><td>Chimaltenango, 4400ft: Municipio de Yepocapa, Finca Recreo</td><td>14.466700</td><td>-90.983300</td></tr><tr><td>Campo Los Guacamayos, Biotopo Laguna Del Tigre, 40 km N</td><td>17.580039</td><td>-90.811972</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Guyo</td><td>16.900561</td><td>-90.293000</td></tr><tr><td>Baramita, North Side Airstrip</td><td>7.371400</td><td>-60.495800</td></tr><tr><td>Kumaka</td><td>7.650000</td><td>-58.950000</td></tr><tr><td>Mount Roraima, Ridge Camp</td><td>5.203125</td><td>-60.736975</td></tr><tr><td>Shanklands</td><td>6.483333</td><td>-58.583333</td></tr><tr><td>Sandstone, Iwokrama Forest</td><td>4.383740</td><td>-58.921220</td></tr><tr><td>Surama Lake</td><td>4.150980</td><td>-59.038460</td></tr><tr><td>Dubulay Ranch</td><td>5.681944</td><td>-57.860556</td></tr><tr><td>Dubulay Ranch, Region 10, Subregion 2</td><td>5.666667</td><td>-57.850000</td></tr><tr><td>Tropenbos, 20 km SSE of Mabura Hill</td><td>5.133306</td><td>-58.724472</td></tr><tr><td>Tropenbos, 20 km SSE of Mabura Hill</td><td>5.133306</td><td>-58.724472</td></tr><tr><td>Rupununi Savanna</td><td>2.999722</td><td>-59.499722</td></tr><tr><td>1 km upstream From Charwir River Mouth, Rupununi River</td><td>2.486389</td><td>-59.476389</td></tr><tr><td>Dadanawa, 15 Mi E, Weramna-Boco, Savannah Creek</td><td>2.824167</td><td>-59.525833</td></tr><tr><td>Dadanawa, 15 Mi ENE, Comiwari Wau</td><td>2.824167</td><td>-59.525833</td></tr><tr><td>Dadanawa, 15 Mi ENE, Comiwari Wau</td><td>2.824167</td><td>-59.525833</td></tr><tr><td>Dadanawa, 30 Mi ENE, Caribuan Forest</td><td>2.824167</td><td>-59.525833</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr></table>
<table><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.711836</td><td>-88.623428</td></tr><tr><td>Las Flores</td><td>14.300000</td><td>-87.833333</td></tr><tr><td>44 km S of Constitucion, 44 km S, 70 km E of Escarcega</td><td>18.250000</td><td>-90.066667</td></tr><tr><td>4 miles west of San Pedro Tapanatepec, Ostuta River</td><td>16.371111</td><td>-94.193056</td></tr><tr><td>6 miles north of Matias Romero</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>6 miles north of Matias Romero</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>6 miles north of Matias Romero</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>6 miles north of Matias Romero</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>6 miles north of Matias Romero</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>Matias Romero, 17 Mi N</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>Matias Romero, 17 Mi N</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>Matias Romero, 17 Mi N</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>Matias Romero, 17 Mi N</td><td>16.878889</td><td>-95.039444</td></tr><tr><td>Palomares, 11.7 Mi S</td><td>17.138333</td><td>-95.062778</td></tr><tr><td>Isla Colon, La Gruta</td><td>9.391167</td><td>-82.277333</td></tr><tr><td>Isla Colon, La Gruta</td><td>9.391167</td><td>-82.277333</td></tr><tr><td>Parque Nacional Soberania</td><td>9.074333</td><td>-79.659778</td></tr><tr><td>Sherman</td><td>9.365000</td><td>-79.958611</td></tr><tr><td>Meteti, Filo del Tallo, Net 2</td><td>8.465395</td><td>-77.992463</td></tr><tr><td>Meteti, Filo del Tallo, Net 3</td><td>8.465395</td><td>-77.992463</td></tr><tr><td>Parque Nacional Darien, Rancho Frio, ANAM Ranger Station,</td><td>8.016700</td><td>-77.729460</td></tr><tr><td>Parque Nacional Darien, Rancho Frio, ANAM Ranger Station,</td><td>8.016700</td><td>-77.729460</td></tr><tr><td>Tacarcuna, Village Camp</td><td>8.165417</td><td>-77.295806</td></tr><tr><td>Tacarcuna, Village Camp</td><td>8.165417</td><td>-77.295806</td></tr><tr><td>Tacarcuna, Village Camp</td><td>8.165417</td><td>-77.295806</td></tr><tr><td>Tacarcuna, Village Camp</td><td>8.165417</td><td>-77.295806</td></tr><tr><td>Tacarcuna, Village Camp</td><td>8.165417</td><td>-77.295806</td></tr><tr><td>La Jagua</td><td>7.866667</td><td>-80.350000</td></tr><tr><td>Summit</td><td>9.066667</td><td>-79.650000</td></tr><tr><td>40 miles east of Panama City, La Cagna River</td><td>8.910111</td><td>-78.846861</td></tr><tr><td>Chilibrillo River</td><td>9.171977</td><td>-79.623222</td></tr><tr><td>Nuri [Montijo District]</td><td>7.979194</td><td>-81.042556</td></tr><tr><td>Chepo Road</td><td>9.167778</td><td>-79.093611</td></tr><tr><td>Chepo Road</td><td>9.167778</td><td>-79.093611</td></tr><tr><td>Chepo Road</td><td>9.167778</td><td>-79.093611</td></tr><tr><td>Chepo Road</td><td>9.167778</td><td>-79.093611</td></tr></table>
<table><tr><td>Chilibrillo Cave</td><td>9.171977</td><td>-79.623222</td></tr><tr><td>Mandinga</td><td>9.450000</td><td>-79.066667</td></tr><tr><td>Mandinga</td><td>9.450000</td><td>-79.066667</td></tr><tr><td>Mandinga</td><td>9.450000</td><td>-79.066667</td></tr><tr><td>Pacora</td><td>9.079444</td><td>-79.290000</td></tr><tr><td>Pacora</td><td>9.079444</td><td>-79.290000</td></tr><tr><td>Pacora</td><td>9.079444</td><td>-79.290000</td></tr><tr><td>Pacora</td><td>9.079444</td><td>-79.290000</td></tr><tr><td>Pacora</td><td>8.966670</td><td>-79.533300</td></tr><tr><td>Panama City, Chepo Rd.</td><td>-9.350000</td><td>-74.510000</td></tr><tr><td>Llullapichis River, tributary of Pachitea River</td><td>-9.370000</td><td>-74.550000</td></tr><tr><td>Nuevo San Juan, Galvez River</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Nuevo San Juan, Galvez River</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Nuevo San Juan, Galvez River</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Nuevo San Juan, Galvez River</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Orosa, Amazon River</td><td>-3.533333</td><td>-72.183333</td></tr><tr><td>Jenaro Herrerra</td><td>-4.905000</td><td>-73.669722</td></tr><tr><td>Sarayacu, Ucayali River</td><td>-6.783333</td><td>-75.116667</td></tr><tr><td>Sarayacu, Ucayali River</td><td>-6.783333</td><td>-75.116667</td></tr><tr><td>Sarayacu, Ucayali River</td><td>-6.783333</td><td>-75.116667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Río Galvez; Nuevo San Juan</td><td>-5.250000</td><td>-73.166667</td></tr><tr><td>Cocha Salvador</td><td>-12.003028</td><td>-71.215694</td></tr><tr><td>Pakitza</td><td>-11.944917</td><td>-71.282972</td></tr><tr><td>Pakitza</td><td>-11.944917</td><td>-71.282972</td></tr><tr><td>Madre de Dios, Manu, 480m: Maskoitania, 13.4 km NNW Ata</td><td>-12.771600</td><td>-71.385400</td></tr><tr><td>Madre de Dios, Manu, 480m: Maskoitania, 13.4 km NNW Ata</td><td>-12.771700</td><td>-71.385500</td></tr><tr><td>San Juan</td><td>-10.371111</td><td>-76.198056</td></tr><tr><td>San Juan</td><td>-10.371111</td><td>-76.198056</td></tr><tr><td>Brownsberg Nature Park</td><td>4.946690</td><td>-55.183450</td></tr><tr><td>Brownsberg Nature Park</td><td>4.946690</td><td>-55.183450</td></tr><tr><td>Sabajo, Transect AF1</td><td>5.093790</td><td>-54.847260</td></tr><tr><td>Bakhuis</td><td>4.740597</td><td>-56.801583</td></tr><tr><td>Kasikasima</td><td>2.977410</td><td>-55.384790</td></tr><tr><td>Merian Site 3</td><td>5.103570</td><td>-54.516580</td></tr><tr><td>Upper Palumeu River</td><td>2.477050</td><td>-55.629410</td></tr><tr><td>Upper Palumeu River</td><td>2.477050</td><td>-55.629410</td></tr><tr><td>Cap-de-Ville</td><td>10.152500</td><td>-61.707500</td></tr><tr><td>Fyzabad</td><td>10.181111</td><td>-61.545833</td></tr><tr><td>Parry Lands, Lot 10</td><td>10.182222</td><td>-61.620833</td></tr></table>
<table><tr><td>Penal Rock Road</td><td>10.133361</td><td>-61.435972</td></tr><tr><td>Edwards Trace</td><td>10.116194</td><td>-61.196833</td></tr><tr><td>Pozon, 50 km NE of Puerto Ayacucho</td><td>6.050000</td><td>-67.416667</td></tr><tr><td>Rio Mavaca, 108 Km SSE Esmeralda</td><td>2.250000</td><td>-65.280000</td></tr><tr><td>Rio Mavaca, 108 Km SSE Esmeralda</td><td>2.250000</td><td>-65.280000</td></tr><tr><td>San Juan, 163 Km ESE Pto. Ayacucho, Rio Manapiare</td><td>5.300000</td><td>-66.220000</td></tr><tr><td>San Juan, 163 Km ESE Pto. Ayacucho, Rio Manapiare</td><td>5.300000</td><td>-66.220000</td></tr><tr><td>Hato Cariben, 32 Km NE Pto. Paez, La Villa</td><td>6.550000</td><td>-67.220000</td></tr><tr><td>Hato Cariben, 32 Km NE Pto. Paez, La Villa</td><td>6.550000</td><td>-67.220000</td></tr><tr><td>Rancho Grande</td><td>10.216667</td><td>-67.416667</td></tr><tr><td>Rancho Grande</td><td>10.216667</td><td>-67.416667</td></tr><tr><td>Rancho Grande</td><td>10.216667</td><td>-67.416667</td></tr><tr><td>Maracay, 4 Km NW, El Limon</td><td>10.283333</td><td>-67.600000</td></tr><tr><td>3 km E of Puerto Cabello Del Caura</td><td>7.166667</td><td>-64.983333</td></tr><tr><td>Hata La Cerbatana, 20 km S of Guaniamo</td><td>6.433333</td><td>-66.116667</td></tr><tr><td>Hato La Florida, 35 km Ese of Caicara</td><td>7.566667</td><td>-65.866667</td></tr><tr><td>Hato San Jose, 20 Km W La Paragua</td><td>6.820000</td><td>-63.480000</td></tr><tr><td>Hato San Jose, 20 Km W La Paragua</td><td>6.820000</td><td>-63.480000</td></tr><tr><td>Icabaru, 45 Km NE Icabaru, Santa Lucia De Surukun</td><td>4.550000</td><td>-61.420000</td></tr><tr><td>km. 125, 85 Km SSE El Dorado</td><td>5.980000</td><td>-61.430000</td></tr><tr><td>km. 125, 85 Km SSE El Dorado</td><td>5.980000</td><td>-61.430000</td></tr><tr><td>San Esteban</td><td>10.433333</td><td>-68.016667</td></tr><tr><td>San Esteban</td><td>11.233333</td><td>-74.133333</td></tr><tr><td>San Esteban</td><td>10.433333</td><td>-68.016667</td></tr><tr><td>Riecito, 30 Km S Mirimiri</td><td>10.900000</td><td>-68.770000</td></tr><tr><td>Urama, 19 Km NW Urama, Km 40</td><td>10.620000</td><td>-68.400000</td></tr><tr><td>Urama, 19 Km NW Urama, Km 40</td><td>10.620000</td><td>-68.400000</td></tr><tr><td>Embalse De Guarico, 10 Km N Calabozo</td><td>9.020000</td><td>-67.430000</td></tr><tr><td>Embalse De Guarico, 10 Km N Calabozo</td><td>9.020000</td><td>-67.430000</td></tr><tr><td>Embalse De Guarico, 10 Km N Calabozo</td><td>9.020000</td><td>-67.430000</td></tr><tr><td>Estacion Biologica De Los Llanos, 14 Km SE Calabozo, Nr. Rio</td><td>8.820000</td><td>-67.370000</td></tr><tr><td>Estacion Biologica De Los Llanos, 14 Km SE Calabozo, Nr. Rio</td><td>8.820000</td><td>-67.370000</td></tr><tr><td>Rio Orituco, 10 km W of Chaguaramas</td><td>9.400000</td><td>-66.466667</td></tr><tr><td>San Jose De Tiznados, 52 Km NNW Calabozo</td><td>9.380000</td><td>-67.570000</td></tr><tr><td>Valera, 23 Km NNW Valera, Rio Motatan</td><td>9.530000</td><td>-70.670000</td></tr><tr><td>Valera, 23 Km NNW Valera, Rio Motatan</td><td>9.530000</td><td>-70.670000</td></tr><tr><td>El Rosario, 60 Km WNW Encontrados, Boca Del Rio De Oro</td><td>9.120000</td><td>-72.770000</td></tr></table>