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The Koepcke’s spear-nosed bat, Gardnerycteris koepckeae (Gardner and

Patton, 1972) (Chiroptera: Phyllostomidae), is not endemic to Peru: first
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Article in Mammalia · September 2020

DOI: 10.1515/mammalia-2019-0107

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Darwin M. Morales-Martínez*, Héctor E. Ramírez-Chaves, Javier E. Colmenares-Pinzón
and Luis G. Gómez
The Koepcke’s spear-nosed bat, Gardnerycteris koepckeae
(Gardner and Patton, 1972) (Chiroptera: Phyllostomidae),
is not endemic to Peru: first record from the Amazon foothills
of Colombia
https://doi.org/10.1515/mammalia-2019-0107
Received August 28, 2019; accepted December 17, 2019
Abstract: We report the first record of the rare Koepcke’s
spear-nosed bat Gardnerycteris koepckeae collected out-
side of Peru, based on morphological and cytochrome b
data. The species was found at the National Natural Park
Alto Fragua Indi-Wasi, Department of Caquetá, Colombia.
This record extended the species distribution 1400 km
north from the northernmost known locality in Huan-
huachayo, Ayacucho, Peru. This finding suggests that G.
koepckeae is distributed over middle elevations along the
eastern slope of the Andean-Amazonia foothills in Peru
and Colombia, similar to other Phyllostomid bats, such
as Vampyressa melissa and Lonchophylla handleyi. We
reviewed the threatened category of G. koepckeae, cur-
rently categorized as Data Deficient (DD), according to the
International Union for Conservation of Nature (IUCN),
but considered as Endangered (EN) in recent papers.
Based on this new record, we suggest that the species
should be included in future assessments as Near Threat-
ened (NT) or Vulnerable (VU) based on its wider distribu-
tion along the Andean-Amazonian foothills of Colombia,
Ecuador and Peru, and probably the Orinoco slope of the
Andes of Colombia and Venezuela.
*Corresponding author: Darwin M. Morales-Martínez, Grupo
Biodiversidad y Conservación Genética, Instituto de Genética,
Universidad Nacional de Colombia and Grupo de Fauna, Programa
de Ecosistemas y Recursos Naturales, Instituto Amazónico de
Investigaciones Científicas SINCHI, Calle 20 # 5-44, Bogota 111321,
Colombia, e-mail: dmmoralesm@unal.edu.co. https://orcid.
org/0000-0001-5786-4107
Héctor E. Ramírez-Chaves: Departamento de Ciencias Biológicas,
Facultad de Ciencias Exactas y Naturales, and Centro de Museos,
Museo de Historia Natural, Universidad de Caldas, Calle 65 # 26-10,
Manizales, Colombia. https://orcid.org/0000-0002-2454-9482
Javier E. Colmenares-Pinzón: Grupo de Estudios en Biodiversidad,
Universidad Industrial de Santander, Carrera 27 # 9, Bucaramanga,
Colombia
Luis G. Gómez: Departamento de Biología, Grupo GECO, Universidad
del Cauca, Popayán, Colombia
Mammalia 2020; aop
Keywords: Andes; distribution; endemism; Mimon;
neotropics.
Introduction
The genus of Neotropical bats Gardnerycteris Hurtado and
Pacheco, 2014 was recently erected based on morphologi-
cal and molecular evidence, re-organizing its previous
paraphyletic position within the genus Mimon sensu lato
(Hurtado and Pacheco 2014, Hurtado and D’Elía 2018).
Gardnerycteris is a well-supported monophyletic group
and, as currently understood, includes three species:
Gardnerycteris crenulatum (É. Geoffroy St.-Hilaire, 1803)
from the eastern lowlands to the Andes, Gardnerycteris
keenani (Handley, 1960) from the western lowlands to the
Andes, and Gardnerycteris koepckeae (Gardner and Patton,
1972) from medium elevations in the Andes cordillera of
Peru (Hurtado and D’Elía 2018). Out of these, G. koepckeae
is the rarest species, because it is only represented by four
specimens from two localities from the eastern slopes of
the Peruvian Andes, at an elevational range between 1600
and 1900 m (Hurtado et al. 2014). Gardnerycteris koepck-
eae is listed as Data Deficient (DD) in the Red List of the
International Union for Conservation of Nature (IUCN)
because there is almost no information about its extent of
occurrence, population trends, and threats and ecological
requirements (Velazco and Aguirre 2019). In this paper,
we present the first record of this rare bat from Colombia,
extending its distribution 1400 km north from the previ-
ously known northernmost locality, and we provide com-
ments on the species status for future assessments.
Materials and methods
A single specimen allocated to Gardnerycteris crenulatum
here was captured using mist-nets in the understory of a
continuous Andean forest on March 15, 2012, at Belén de
los Andaquíes, La Quisayá stream, National Natural Park
Alto Fragua Indi Wasi, Alto Río Pescado, Department of
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2 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia

Caquetá (1°34′48″N; 75°56′2.85″W, 1080 m; Figure 1) in
southwestern Colombia. This area comprises a mature
sub-Andean forest, dominated by palms up to 30 m high
with epiphytes (Rodríguez et al. 2006) and mixed with
patches of low secondary vegetation. The weather is
relatively warm throughout the year, with temperatures
ranging from 25 to 30°C and precipitation between 4000
and 5000 mm (Robertson and Castiblanco 2005, Gómez
et al. 2016). The collected specimen was preserved as skin
and skull and deposited in the mammal collection of the
Museo de Historia Natural, Universidad del Cauca, under
catalog number MHNUC 2010.
Morphological analysis
To corroborate the taxonomic identity of this specimen,
we examined and compared the specimen (MHNUC
2010) with all the specimens of the genus Gardneryct-
eris deposited in the following collections in Colombia

Figure 1: Distribution of Gardnerycteris koepckeae.

(1) First record from Colombia: Belén de Los Andaquíes, La Quizayá, National Natural Park Alto Fragua Indi Wasi, Alto río Pescado,
Department of Caquetá (1°34′48″N; 75°56′2.85″W; 1080 m). (2) Type locality: Huanhuachayo, Ayacucho Department (12°44′00″S,
73°47′00″W; 1660 m). (3) Podocarpus, Santuario Nacional Pampa Hermosa, Chanchamayo District, Chanchamayo Province, Junin
Department (10°59′49.2″S, 75°25′57.1″W; 1890 m).

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 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia
(Appendix 1): Colección de Mamíferos “Alberto Cadena
García” at the Instituto de Ciencias Naturales, Universi-
dad Nacional de Colombia (ICN), Colección de Mamíferos
del Museo de Historia Natural de la Universidad del
Cauca (MHNUC), and the Centro de Museos, Museo de
Historia Natural de la Universidad de Caldas (MHN-UCa).
We carried out species identification using the diagnostic
characters provided for the species of Gardnerycteris by
Hurtado and Pacheco (2014), Hurtado et al. (2014), and
Hurtado and D’Elía (2018).
Additionally, we recorded all external and cranioden-
tal measurements using a digital caliper to the nearest
0.1 mm following Hurtado et al. (2014). We took the
external measurements (except forearm length) from the
specimen’s tags. We performed a principal component
(PC) analysis on 16 craniodental measurements using the
statistical software PAST version 2.2 (Hammer et al. 2001)
including 32 individuals, 26 Gardnerycteris crenulatum,
six Gardnerycteris keenani, and one Gardnerycteris koep-
ckeae. The analysis was performed using the covariance
matrix to preserve the information about relative scale
among variables.
Molecular analysis
To further ascertain the identification of the specimen
MHNUC 2010, we carried out a molecular characterization
using the mitochondrial marker, the cytochrome b (Cyt
b). We selected this marker because of its relevance for
clarifying the taxonomic status of other mammals whose
taxonomy had been predominantly studied based only
on morphology and/or karyological data (Gutiérrez et al.
2017), and because there are available sequences of the
three species of Gardnerycteris in GenBank. We extracted
total DNA from a small piece of bone clipped from the
forearm bone. We used the classic phenol/chloroform
protocol (Sambrook et al. 1989). Then, 923 base pairs of
the Cyt b were polymerase chain reaction (PCR)-amplified
using the primers in Bradley and Baker (2001): glo7L
(5′-CAY CGT TGT ATT TCA ACT RTA AGA AC-3′) and glo6H
(5′-CGG TGT AAT GRA TAT ACT ACA TRG-3′). We performed
a PCR in a Multigene Thermal Cycler TC9600-G (Labnet
International, Inc., Edison, NJ, USA). The program con-
sisted of an initial denaturing step at 94 for 2 min followed
by 35 cycles of denaturation at 94 for 45 s, annealing at
50.5 for 1 min and extension at 72°C for 1 min, and a final
extension step at 72°C for 15 min. The PCR was carried out
in a total volume of 30 μl with approximately 1.5 μl of DNA,
1X buffer (with 2.0 mm of MgCl2), 0.6 mm of deoxyribonu-
cleotide triphosphates (dNTPs), 1.5 μm of each primer,
3
and 0.15 U/μl of DreamTaq DNA Polymerase (Thermo Sci-
entific, Waltham, MA, USA). We carried out purification
and sequencing of both strands with the amplification
primers on an ABI 3500 sequencer (Applied Biosystems,
Waltham, MA, USA) at the “Servicio de Secuenciación y
Análisis ­Molecular SSiGMol” at the Universidad Nacional
de Colombia, Colombia, Bogotá. Sequences from each
primer were checked for quality and assembled using the
program Geneious Prime (Biomatters Limited, Auckland,
New Zealand). The resulting sequence was translated to
protein, also in Geneious Prime (2019.2.1), to avoid prema-
ture stop codons and to verify its correct edition. Posteri-
orly, we submitted it to GenBank where it can be found
under the accession number MN327642.
We analyzed our sequence from specimen MHNUC
2010 in a phylogenetic framework in order to assess its
affinities and relationships with other Gardnerycteris
species. We used a Bayesian inference approach on
a matrix with 38 terminals that included the same 36
Cyt b sequences reported by Hurtado and D’Elía (2018),
plus the sequence from MHNUC 2010 and a second new
sequence from a specimen of Gardnerycteris crenulatum
collected in the middle Magdalena Valley in ­Colombia.
We aligned all the sequences using the default para-
meters of the Multiple Sequence Comparison by Log-
Expectation (MUSCLE) algorithm as implemented in
Geneious Prime. We conducted Bayesian analysis in
MrBayes 3.2.6 (Ronquist et al. 2012), specifying the
GTR + I + Γ as the best fitted nucleotide substitution
model, based on the Akaike information criterion (AIC)
implemented in jModelTest 2.1.10 (Darriba et al. 2012). To
ensure convergence, we ran two independent replicates
of the Metropolis-coupled chain Monte Carlo analysis for
100,000,000 generations with trees being sampled every
1000 generations. We inspected stationarity and conver-
gence in the program Tracer 1.6 (Rambaut et al. 2014) by
plotting likelihood values per generation. We discarded
25% of the samples in each run as burn-in and we com-
bined the remaining samples (75,000) to estimate tree
topology, the mean likelihood, and posterior probabili-
ties. We considered that a node received strong (signifi-
cant) support when its posterior probability was >0.95%
(Gutiérrez et al. 2017).
To improve our comparative framework, we used the
program MEGA 7 (Kumar et al. 2015) to calculate average
uncorrected p-distances and model-corrected Kimura
2-parameter (K2P) distances in the Cyt b matrix. The per-
centage of genetic divergence was computed as genetic
distance × 100. We contrasted the values obtained with
those reported for bats by Bradley and Baker (2001), and
Hurtado and D’Elía (2018) for Gardnerycteris.
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4 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia

Results et al. 2014): dorsal fur is unicolored without a dorsal

The new record is an adult male (MHNUC 2010) that
matches morphologically the description and re-descrip-
tion of Gardnerycteris koepckeae, including the following
diagnostic characters (Gardner and Patton 1972, Hurtado
white stripe, ventral coloration is bicolored; the length
of the noseleaf is twice its width; the forearm is bare;
the insertion of the plagiopatagium is at the tarsus level
on the V metacarpal; cranially, nasal depression has a
V-shape compared with other Gardnerycteris; the hard

Figure 2: Morphological and cranial characters of Gardnerycteris koepckeae from Colombia (MHNUC 2010).
Scale bar 10 mm. (A) skin in dorsal view, (B) skin in ventral view, (C) insertion of the plagiopatagium at the tarsus level, (D) Dorsal (top),
ventral (middle) and lateral (bottom) views of the skull.

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 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia 5

palate passes the middle of the mesopterygoid fossa and
has a V-shape; and M1 and M2 have a cleft between the
protocone and the hypocone (Figure 2). Measurements
are in the ranges of the specimens reported by Hurtado
et al. (2014) except for the postorbital width, zygomatic
width, and the palatal length that are slightly lower, and
the palatal width that is slightly larger (Table 1). PC 1 and
2 accounted for 74.4% of the total variance (PC 1 = 60.8%
and PC 2 = 10.6%), but only PC 2 component show differ-
ences between species and was correlated with the size of
the auditory bulla (loading = 0.76) and the size of the skull
(loading = −0.42). Our specimen is isolated from Gard-
nerycteris crenulatum and Gardnerycteris keenani groups
in the morphometric space (Figure 3).
Our phylogenetic analysis of the Cyt b matrix yielded
a topology that is almost identical to that reported by
Hurtado and D’Elía (2018) for the same marker, with only
subtle differences in the support values at some nodes
(Figure 4). The MHNUC 2010 Cyt b sequence of the speci-
men identified as Gardnerycteris koepckeae using morpho-
logical characters clusters with all the sequences of Cyt b of
all three Gardnerycteris species from GenBank included in
the analysis. Thus, the monophyly of the genus is strongly
supported (PP = 1.00) as the sister group of Phyllostomus
(as in the topology presented by Hurtado and D’Elía 2018).
Within Gardnerycteris, the haplotype of MHNUC 2010
is sister to the only other available Cyt b haplotype of G.
koepckeae (PP = 1.00). This close relationship confirms
the taxonomic identity of MHNUC 2010 as G. koepckeae.
Our analysis recovered the same pattern of relationships
between species of Gardnerycteris suggested by Hurtado
and D’Elía (2018), thereby supporting the hypothesis that
Gardnerycteris keenani is the most basal species in the
genus, and that it is sister to a monophyletic group con-
formed by Gardnerycteris crenulatum and G. koepckeae.
Cyt b genetic distances also provide evidence that
specimen MHNUC 2010 is Gardnerycteris koepckeae. The
percentage of divergence between its haplotype and the
only other available haplotype of the species is 2.8 ± 0.5%
(p-distance) or 2.9% (K2P), whereas the percentages
between MHNUC 2010 and the remaining Gardnerycteris
haplotypes are in the range of 9.4 ± 0.8% (from Gard-
nerycteris crenulatum) to 11.1 ± 0.9% (from Gardnerycteris
keenani), considering only p-distances (Table 2).
Discussion
Our results imply that the morphological characters are
reliable for an accurate identification of this species. Addi-
tionally, the value of the genetic distance of Cyt b gene (2.9%
K2P) between two specimens of Gardnerycteris koepckeae
was consistent with the typical mean intraspecific diver-
gence reported by Bradley and Baker (2001) for bats, and a
value of 2.8% (p-distance) was within the range reported by
Velazco et al. (2010), Velazco and Simmons (2011), Velazco
and Cadenillas (2011), and Martínez-Cerón et al. (2019)
for phyllostomid species of Platyrrhinus, Vampyrodes,

Table 1: Comparative measurements of species of Gardnerycteris.

G. koepckeae G. crenulatum G. keenani

Hurtado et al. 2014 MHNUC 2010

Total length 79.00 ± 1.41 (2) 76.82 79.72 ± 6.64 (18) 84.00 ± 3.61 (3)
Tail length 19.50 ± 4.95 (2) 21.09 23.21 ± 2.86 (19) 23.25 ± 5.12 (4)
Hindfoot length 10.00 ± 1.41 (2) 10.57 10.5 ± 0.90 (12) 10.75 ± 0.96 (4)
Ear length 22.50 ± 0.71 (2) 22.23 23.47 ± 1.98 (19) 22.3 ± 1.92 (5)
Forearm length 47.70 ± 0.42 (2) 48.09 47.64 ± 1.79 (19) 48.47 ± 1.9 (5)
Greatest length of skull 21.11 ± 0.74 (3) 20.83 21.86 ± 0.55 (26) 21.79 ± 0.66 (5)
Condylo-incisive length 18.79 ± 0.21 (3) 18.71 19.25 ± 0.47 (26) 19.26 ± 0.36 (5)
Postorbital width 4.14 ± 0.06 (3) 4.02 4.23 ± 0.14 (26) 4.14 ± 0.15 (5)
Zygomatic width 11.67 ± 0.24 (3) 11.25 11.83 ± 0.32 (25) 11.71 ± 0.1 (2)
Braincase width 8.41 ± 0.06 (2) 8.35 8.19 ± 0.21 (26) 8.06 ± 0.22 (5)
Palatal width at canines 5.01 ± 0.01 (3) 5.04 5.31 ± 0.16 (26) 5.12 ± 0.13 (4)
Mastoid width 9.92 ± 0.86 (3) 10.48 11.26 ± 0.37 (26) 11.27 ± 0.16 (5)
Palatal length 8.44 ± 0.33 (3) 7.62 8.46 ± 0.43 (26) 8.48 ± 0.16 (4)
Width at M1-M1 7.43 ± 0.18 (2) 7.33 7.59 ± 0.17 (26) 7.41 ± 0.29 (5)
Width at M2-M2 8.04 ± 0.19 (3) 8.17 8.15 ± 0.17 (26) 8.05 ± 0.33 (5)
Auditory bullas width 2.36 ± 0.19 (2) 2.3 3.57 ± 0.26 (25) 3.60 ± 0.14 (5)
Maxillary toothrow length 7.71 ± 0.36 (3) 7.8 7.78 ± 0.21 (26) 7.68 ± 0.26 (5)
Molariform toothrow length 6.64 ± 0.74 (3) 6.33 6.42 ± 0.20 (26) 6.63 ± 0.19 (5)

Mean ± standard deviations (n).

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 6 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia

1.0

Gardnerycteris keenani
0.5
Gardnerycteris crenulatum
Component 2 (10.58%)

–3.0 –2.5 –2.0 –1.5 –1.0 –0.5 0.5 1.0 1.5

–0.5

–1.0

–1.5
Garnerycteris koepckeae

–2.0
Component 1 (60.85%)

Figure 3: Scatter plot of the first and second principal component scores of a principal component analysis based on 16 cranial and
mandibular measurements from three species of Gardnerycteris including G. crenulatum, G. keenani, and G. koepckeae.

Figure 4: Cyt b tree recovered from the Bayesian inference analysis.

Support statistics (posterior probabilities) are indicated at each resolved node. Gray box contains the two haplotypes of Gardnerycteris
koepckeae. Each Gardnerycteris terminal is identified by country of origin and next two largest political units.

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 D.M. Morales-Martínez et al.: First record of Gardnerycteris koepckeae from Colombia
Table 2: Cyt b sequence divergence (%) between and within
Gardnerycteris species.
1 2
1-Gardnerycteris koepckeae 2.8 12.2
± 0.5 ± 1.1
2-Gardnerycteris keenani 11.1 0.3
± 0.9 ± 0.1
3-Gardnerycteris crenulatum 9.4 10.6
± 0.8 ± 0.9
Uncorrected pairwise interspecific divergences are shown below
the diagonal, the K2P model corrected above the diagonal, and
intraspecific uncorrected p-distances in the diagonal (in bold). Numbers
below each divergence percentage represent standard errors.
Lophostoma, and Phylloderma, respectively. Furthermore,
K2P-based divergences between the two G. koepckeae hap-
lotypes and the remaining Gardnerycteris haplotypes were
consistent with the mean interspecific divergences for
bats, also reported by Bradley and Baker (2001) and with
the p-distance-based interspecific divergences reported by
Hurtado and D’Elía (2018) for Gardnerycteris.
The record of Gardnerycteris koepckeae described
here increases the latitudinal distribution of the species
by 1400 km and represents its lowest elevational collect-
ing site. It also constitutes the third and northernmost
­published locality for this rare bat (Hurtado and D’Elía,
2018), and confirms its presence outside of Peru, increas-
ing the number of mammal species known in Colombia
from 528 to 529 (Ramírez-Chaves et al. 2019). The vast
gap between the Colombian and Peruvian records of the
species clearly suggests its presence in Ecuador, which
surely will be confirmed in the future. In the light of this
new collecting locality, it is evident that G. koepckeae
occupies a wider area than previously thought, as well as
a wider elevational gradient. The two previously known
localities suggested that G. koepckeae was an endemic
montane-highland species, possibly restricted to Andean
ecosystems in central Peru (Hurtado and D’Elía 2018).
However, the lower elevational record from Colom-
bia suggests its presence in the A ­ndean-Amazonian
foothills, within the Amazonian biogeographic prov-
ince (­Hernández-Camacho et al. 1992). This province
encompasses the foothills of Putumayo and Caquetá in
Colombia and extends to the Ecuadorian and northern
Peruvian foothills, an area that is considered to be of
high diversity and endemism in Colombia and the Neo-
tropics (­Hernández-Camacho et al. 1992). The distribu-
tion of G. koepckeae resembles that of other phyllostomid
bats, such as Vampyressa melissa (Tavares et al. 2014)
7
and Lonchophylla handleyi Hill, 1980 (Mantilla-Meluk
et al. 2010), reflecting the peculiarity of the Andean-
Amazonian foothills. The low divergence between the
3
Colombian and Peruvian Cyt b haplotypes of the species
10.3 also constitutes an evidence of gene flow throughout the
± 0.9 species distribution, probably promoted by its historical
11.7
ecological homogeneity. Furthermore, having substan-
± 1.0
5.6 tially increased our knowledge of the distribution of this
± 0.7 species, this record suggests that probably the species has
a wider distribution along the Amazonian and Orinoco
mountain slopes of the Andes associated at middle eleva-
tions (1000–1900 m), like V. melissa (Tavares et al. 2014),
L. handleyi (Mantilla-Meluk et al. 2010), and Lonchophylla
orienticollina Dávalos and Corthals, 2008.
Conservation
Gardnerycteris koepckeae has been categorized as DD in
the IUCN Red List (Velazco and Aguirre 2019). Hurtado
et al. (2014) suggested a revision of the conservation
status of G. koepckeae as Endangered (EN) according
to the criterion B2abi,v of the IUCN (2012), based on the
Peruvian endemism of G. koepckeae, the potential loss of
its habitat, and the rarity of records. The new locality in
Colombia shows that the species does not meet all these
criteria. Now G. koepckeae is distributed over more than
2000 km2 including protected areas such as the National
Natural Park Alto Fragua Indi-Wasi in Colombia. However,
the new record comes from areas with high levels of defor-
estation (Amazon) in Colombia (IDEAM 2014). Even if G.
koepckeae seems to be rare (category D according to Yu
and Dobson 2000), it is possible that the rarity is due to
a lack of sampling efforts in the Andean foothills of the
Amazon and Orinoco basin, Colombia (Hernández-Cama-
cho et al. 1992, Salaman and Donegan 2007). In any case,
there is a need for reassessing the species category, which
perhaps would better meet the criteria for “Vulnerable” or
“Near Threatened”, considering that the species is more
widely distributed in South America.
Acknowledgments: We thank Hugo López, (ICN), Catalina
Cárdenas (ICN), Claudia Medina (IAVH), Andrew Craw-
ford (MUA), Hernando Vergara and Pilar Rivas (MHNUC)
for allowing us to study the specimens under their care.
DMM thanks Rendón-Mera and Ramírez-Chaves’ families
for their assistance during his visit to Popayán city. HERC
thanks Vicerrectoría de Investigaciones, Universidad de
Caldas (project 0743919) for support.
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Appendix 1: Revised specimens

Gardnerycteris crenulatum (n = 27): Colombia: Amazonas, Leticia (ICN 17863 , ICN 17864 , ICN 21029 ); Caquetá:
Río Mesay (ICN 14706 ); Casanare: Sabanalarga (ICN 21003 ); Guainía: Puerto Inírida (ICN 13906 ); Meta: Acacías
(ICN 21564 ), Puerto Gaitán (ICN 5904, ICN 18389 ), San Juan de Arama (ICN 10228 , ICN10229 , ICN 10230 , ICN
10231 , ICN 10232 , ICN 10233 , ICN 10234 , ICN 10235 ), Villavicencio (ICN 7667 , ICN 14338 , ICN 14339 , CIN
14340 , ICN 21091 , ICN 21666 , ICN 22004 ); Putumayo: Puerto Leguízamo (ICN 22004 ); Vaupés: La Libertad
(ICN 12728 ).
Gardnerycteris keenani (n = 11): Caldas: Manizales (MHN-UCa 272-275); Chinchiná (MHN-UCa 2663); Córdoba: Pueblo
Nuevo (ICN 17240 ); La Guajira: Albania (ICN 18630 ), Barrancas (ICN 14754 , ICN 14755 ); Magdalena: Sitio Nuevo
(ICN 15456 ); Santander: Cimitarra (MHN-UCa 1020); Valle del Cauca: Calima (ICN 8889 ).
Gardnerycteris koepckeae (n = 1); Caquetá: San José del Fragua (MHNUC 2010 ).

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