Research Article |
Corresponding author: Angie P. Penagos ( rdangiepenagos@gmail.com ) Academic editor: Ana Maria Leal-Zanchet
© 2020 Diego A. Esquivel, Carlos Aya-Cuero, Angie P. Penagos, Julio Chacón-Pacheco, Carlos J. Agámez-López, Andrés Vargas Ochoa, Héctor E. Ramírez-Chaves, David Bennett.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Esquivel DA, Aya-Cuero C, Penagos AP, Chacón-Pacheco J, Agámez-López CJ, Ochoa AV, Ramírez-Chaves HE, Bennett D (2020) Updating the distribution of Vampyrum spectrum (Chiroptera, Phyllostomidae) in Colombia: new localities, potential distribution and notes on its conservation. Neotropical Biology and Conservation 15(4): 689-709. https://doi.org/10.3897/neotropical.15.e58383
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The Spectral Bat (Vampyrum spectrum) is classified as Near Threatened by the International Union for Conservation of Nature. In Colombia, it has not been included in any risk assessments, because it is a poorly documented species in the country. Using new occurrence data and ecological niche modelling we evaluate the distribution and analyze the species’ ecological and conservation needs in Colombia. With the new records, V. spectrum shows a distribution over six biogeographic provinces in the country, along an elevational range between 96 and 1750 m. The Spectral Bat has been reported in a wide variety of ecosystems from dry forest and fragmented landscapes, to forested areas. Predictive models suggest a wider presence of this species in the west of the country, center, and southern Andes. The limited number of specimens limits our attempt to evaluate morphometric variation among Colombian populations; however, we detect a variation in the mean value of the forearm and ear length regarding northeast South American populations. Areas with the greatest potential distribution detected in this study could be prioritized in conservation strategies, and make plans accordingly for this and other species that coexist in its range. Finally, we highlight the low representativeness of Protected Areas for the Spectral Bat and add some ecological notes about this rare species.
Carnivorous bat, ecological niche modelling, Phyllostominae, Protected Areas, rare species, Spectral Bat
The Spectral Bat Vampyrum spectrum (Linnaeus, 1758) is the largest bat species in the Neotropical region. It has a wide distribution ranging from southern Mexico to northern and central Bolivia (
Globally, the Spectral Bat is listed as Near Threatened (NT) according to the IUCN Red List (
Since then, the presence of the species has been confirmed in different geographic regions of Colombia including: i) The Andean Region: departments of Antioquia (
Correlative environmental niche modelling (ENM) attempts to model a species’ relationship with environmental variables and predict habitat suitability for that species based on known observations of its presence. Although ENM techniques are inherently limited by imperfect knowledge, they have nonetheless proven extremely useful in assessing the effectiveness of Protected Areas distribution (
In this study we (1) report new localities of V. spectrum from the Andean, Caribbean and Orinoquian regions, (2) update the distribution of the species in Colombia, (3) use the distribution data together with Ecological Niche Modelling (ENM) to generate a map of the potential distribution of the species, and (4) include information on its morphometric variation, conservation status and some ecological notes that contribute to the knowledge of this rare species.
We compiled a dataset of occurrence records of the Spectral Bat V. spectrum throughout its entire distribution in the Neotropics by searching in on-line databases including the Arctos database (
To update the distribution of V. spectrum in Colombia, we also reviewed specimens from biological collections (Appendix
To adequately characterize the V. spectrum’s environmental niche, we took 183 occurrence points from its entire neotropical distribution and thinned them by 5 km to reduce spatial autocorrelation – distance based on home range (
The final ensemble map was reclassified using the binary threshold. This binary map was then further filtered in ArcMap v10.6 to exclude “above threshold” pixels in areas above 2000 m a. s. l (taking into account that 1750 m is the maximum elevation for V. spectrum in Colombia), using the GMTED elevation dataset (
We took external (N = 3) and craniodental (N = 13) measurements using digital calipers of 0.01 mm accuracy, following
In Colombia, Vampyrum spectrum is known from 37 records in 15 departments along six biogeographic provinces: a) Peri-Caribbean Arid Belt, b) Orinoco, c) Guyana, d) Amazonia e) Chocó-Magdalena and f) Nor-Andean. Likewise, the species has been recorded in different ecoregions as Montane forests (19 records), Moist forests (8), Dry forests (4), Llanos (4) and Xeric scrub (2). Based on vouchers, the Spectral Bat is distributed in an elevational range between 96 and 1750 m, with most records (67.5%; N = 25) below 600 m. Of the total, four are new localities, recently obtained in fieldwork activities (Fig.
Records of Vampyrum spectrum in Colombia. * indicates new records and *** indicate the highest record in the country. Acronyms: Colección Teriológica Universidad de Antioquia (
Coordinates (Latitude, Longitude) | Department | Locality | Record Year | Elevation (m.a.s.l) | Reference | Records from museums | |
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1 | -0.6217, -72.384 | Amazonas | Araracuara: Hospital | 1977 | 192 |
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2 | -0.6217, -72.384 | Amazonas | Araracuara: Hospital | 1977 | 192 |
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3 | -0.6217, -72.384 | Amazonas | Colonia de Araracuara | 1961 | 200 |
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4 | -1.3003, -69.564 | Amazonas | Corregimiento La Pedrera; Rio Caquetá, Quebrada El Ayo | 2001 | 96 |
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5 | 7.0833, -75.133 | Antioquia | Anori |
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6* | 7.2307, -76.450 | Antioquia | Corregimiento Pavarando Grande; Vereda Chontadural | 2019 | 163 | This study | CZUC-M 0409 |
7 | 5.7668, -74.974 | Antioquia | Vereda Campo Alegre; Predios de Don Guillermo Garcés | 2016 | 646 |
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8 | 6.9114, -74.657 | Antioquia | Vereda La Cruz; Finca La Brillantina | 2009 | 550 |
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9 | 7.4833, -74.867 | Antioquia | Zaragoza; 25 Km S La Tirana | 1972 | 520 |
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10 | 7.4870, -74.868 | Antioquia | Zaragoza; Providencia | 1972 | 410 |
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11 | 5.4500, -74.667 | Caldas | La Dorada |
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12 | 4.9969, -75.594 | Caldas | Manizales; Reserva Planalto | 2007 | 1350 |
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MHN-UCa 959 |
13 | 5.6689, -74.788 | Caldas | Norcasia; Vereda Quiebra de Roque | 2010 | 250 | MHN-UCa 989 | |
14 | 5.5729, -74.940 | Caldas | Norcasia; Vereda Monte Bello | 2015 | 505 | MHN-UCa 1478 | |
15 | 5.3161, -74.833 | Caldas | Victoria; Carretera Victoria-Mariquita; Vereda El Llano | 1967 | 285 |
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16 | 5.3161, -74.833 | Caldas | Victoria; Carretera Victoria-Mariquita; Vereda El Llano | 1967 | 285 |
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17 | 5.3161, -74.833 | Caldas | Victoria; Carretera Victoria-Mariquita; Vereda El Llano | 1967 | 285 |
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18* | 5.4901, -71.118 | Casanare | Paz de Ariporo; El Desengaño | 2017 | 330 | This study | Photograph by Jhon Amaya |
19 | 0.0667, -72.433 | Caquetá | PNN Serranía del Chiribiquete; Rio Mesay, Puerto Abeja | 2001 | 240 |
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20 | 0.2322, -72.250 | Caquetá | Rio Mesay; Chorro Mazaca | 1995 | 240 |
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21 | 2.2564, -76.869 | Cauca | Coconuco |
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22* | 7.7500, -73.383 | Cesar | San Alberto | 2015 | 125 | This study | Photograph by Luz Myriam Moreno Bejarano |
23 | 4.5167, -73.367 | Cundinamarca | Medina; Granja Experimental Agropecuaria | 1986 | 520 |
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24 | 11.1333, -72.917 | Guajira | Riohacha; Corregimiento Tomarrazon Vereda Colon, Arroyo Los Coquitos | 2011 | 170 |
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25 | 1.8808, -76.270 | Huila | San Agustín | 1952 | 1700 |
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26 | 4.1000, -73.800 | Meta | Acacías; Manzanares 20 Km suroccidente Villavicencio | < 1990 |
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27 | 3.6350, -71.761 | Meta | Puerto Gaitán; vereda Alto de Tillavá, finca Los Cayenos | 2015 | 174 |
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Photography |
28 | 4.0626, -73.515 | Meta | Villavicencio; Hacienda El Hachón Km. 16 carretera Puerto López | 1982 |
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29 | 1.2866, -78.074 | Nariño | Barbacoas; Reserva Natural Rio Ñambi | 2004 | 1450 |
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30*** | 1.5833, -76.967 | Nariño | Ricaurte; Reserva Natural La Planada | 1984 | 1750 |
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31 | 1.6741, -77.302 | Nariño | Taminango; Remolino, El Algodonal | 2013 | 570 |
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32 | 2.5333, -78.267 | Nariño | Tola; Parque Nacional Natural Sanquianga | 2000 |
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33 | 9.5298, -75.352 | Sucre | Coloso; Estación meteorológica primates | 2012 | 300 |
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No voucher |
34* | 4.2900, -75.036 | Tolima | Gualanday | 2015 | 541 | This study | Photograph by Tatiana Toro-Sánchez. |
35 | 4.0167, -74.967 | Tolima | Guamo; Monte Cañada | 1968 | 320 |
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36 | 4.5996, -74.829 | Tolima | Piedras; cuenca Rio Totaré Vereda La Manga de los Rodríguez | 2007 | 270 |
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37 | 4.5000, -69.750 | Vichada | Santa Teresita | 1967 | 162 |
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The first record from Mutatá, Department of Antioquia – north Urabá sub-region, “Vereda” Chontadural (7°13'50.4"N, 76°26'58.8"W; 163 m a. s. l) – is a specimen found as a result of a biological characterization in a requirement for obtaining environmental permits for the development of a mining project (stone), in June 2019. The specimen is an adult male (CZUC-M 0409), found entangled in a wire fencing at ground level on a gutter created for runoff. Because the skin was already totally destroyed, with only the skull preserved, a fracture in the braincase was presumed to be the cause of death. The area containing this specimen was a wooded cover of tropical humid forest (bh-T) crossed by the Caño Bijará, where secondary vegetation prevails with a wide canopy providing spaces for the species’ flight capacity. The second record comes from Paz de Ariporo, Department of Casanare (5°29'24.36"N, 71°7'4.79"W; 330 m a. s. l), and corresponds to an adult male captured as result of a biological characterization during 2017. The specimen was photographed and then released in the same area, which has gallery forests and extensive plains. The third record comes from San Alberto, Department of Cesar (7°45'0"N, 73°22'58.7"W; 125 m a. s. l); it is an individual found in the development of a biological characterization during 2015. The individual corresponds to an adult female recorded in a gallery forest fragment, in a very narrow strip of secondary vegetation on the banks of a stream, near to oil palm crop. The area where this specimen was found is strongly pressured by the oil crop, but there are still some remnants of gallery forest that allow the survival of the species. The fourth record comes from the municipality of Gualanday, Department of Tolima (4°17'24"N, 75°2'9.6"W; 541 m a. s. l), and represents an adult female found as a result of the development of an environmental impact study during 2015. The individual was recorded by Tatiana Toro-Sánchez in a dry tropical forest remnant near water bodies and agricultural crops (Fig.
Our models had a mean TSS (0.49 for BRT, 0.48 for Maxent, 0.47 for RF and 0.46 for SVM) and mean AUC (0.78 for BRT, 0.76 for Maxent, Random Forest and SVM) indicating the models fit well to the data. All individual model runs had values above our minimum accuracy values also. The relative habitat suitability for V. spectrum in Colombia based on our models, and the areas converted into a binary threshold and subsequently filtered to remove areas above 2000 m in elevation, are shown in Fig.
A Potential distribution of Spectral Bat Vampyrum spectrum in Colombia using an ensemble modelling approach with five different algorithms. Blue to red color scales indicate less to greater suitability area for the species. B Ensemble suitability map converted to a binary of above (suitable) and below (not suitable) threshold, manually reclassified to exclude high elevation areas (>2000 m a. s. l). Areas considered suitable were focused in the west and south of the country, although eastern Colombia also had significant areas above the threshold.
Regarding variable importance, although the degree of importance differed between the two importance assessment metrics (TSS/AUC), the order was constant in both. Under both importance assessment methods, forest cover was by far the most important variable, followed by Bio14 (precipitation in the driest month), then slope, and finally Bio05 (maximum temperature of the warmest month) (Table
Variable | AUC metric | Pearson correlation |
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Bio05 | 12.8 | 7.1 |
Bio14 | 40.4 | 19.7 |
EDForest | 53.7 | 29 |
Slope | 17.6 | 7.5 |
The limited number of specimens limits our attempt to evaluate morphometric variation among Colombian populations. In general, the external and craniodental values were similar between specimens from Colombia, Central America and northeast South America (Guyana, French Guyana, Suriname and Trinidad). Despite that, we observed subtle differences in the length of the forearm mean value, and – notably – the ear length that were larger in Colombian populations (Table
External and cranial measurements (in millimeters) of Vampyrum spectrum in Colombia. Mean, range and number individuals are shown. * Data from specimens in museums and literature:
Measurements | Specimens from | ||
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Colombia | Central America* | Northeast South America* | |
FA** | 108.7 (104–115) 16 | 104.8 (95.1–110.4) 9 | 103.9 (100–107.1) 12 |
E** | 41.6 (39–45.7) 18 | 36.0 (30–40) 6 | |
HF | 30.9 (27.6–34.4) 17 | 32.8 (31–35) 7 | 30.0 (29–31) 6 |
GLS | 51.4 (49.29–53.4) 16 | 51.4 (48.1–53.6) 10 | 51.4 (49.2–53.3) 13 |
CIL | 43.1 (41.05–44.51) 14 | 43.2 (41.6–44) 7 | |
CCL | 42.4 (40.49–45.04) 12 | 43.1 (41.8–43.7) 7 | |
ZB | 24.1 (23.43–24.47) 15 | 24.2 (23.1–25.4) 9 | 24.0 (23.2–25.2) 12 |
MTRL | 19.3 (17.11–21.23) 16 | 20.6 (19.7–21.7) 11 | 21.0 (20.6–21.5) 5 |
MANDL | 21.5 (18.59–23.75) 15 | ||
PB | 8.1 (7.65–8.06) 17 | 8.1 (7.8–8.6) 9 | 7.9 (7.3–8.6) 13 |
BB | 16.7 (14.02–19.15) 14 | 16.2 (15.6–18.2) 5 | 15.9 (15.7–16.5) 8 |
BH | 17.7 (15.5–18.37) 13 | ||
PL | 24.6 (23.15–26.8) 15 | 24.8 (23.7–26.1) 7 | |
C-C | 8.7 (7.46–9.25) 14 | ||
SCL | 8.0 (7.43–8.76) 13 | ||
ICL | 8.4 (7.92–9.15) 13 |
V. spectrum has been reported in a wide variety of ecosystems in the country. In the Andean region, most of the reviewed specimens were captured in areas with secondary vegetation and with strong human intervention. Some specimens were found in fragmented forest with the presence of cattle, selective felling and crops, and others have even been found dead resulting from their impact with High Voltage Transmission Lines in Antioquia (Instituto Alexander von Humboldt –
Our model predicted that currently around 13% of total habitat suitability area of Vampyrum spectrum are included within Protected Areas in Colombia. These Protected Areas are mainly concentrated in the Andean, Orinoquia and Amazonia regions. However, at a regional scale, the representativeness of these areas remains very low. PAs barely cover 12.8 and 8.4% of the suitable area in Andean and Orinoquia regions respectively while in the Caribbean region this value is even lower and only reaches 7.6%.
Vampyrum spectrum presents a wide but discontinuous distribution in Colombia, probably related to its low detection. Information about this species is limited and mainly restricted to vouchers’ records in natural history collections. From the first record in 1952 (Museo de Historia Natural de la Universidad del Cauca –
The number of records occurring after 2010 (N = 11), indicate the presence of the species in well-preserved areas in Amazonas, Caquetá, Meta, and Nariño, and its subsistence in patches of forest with strong anthropogenic pressure in the Central Andes. Although efforts are required to determine the state of the populations and confirm their presence in other areas, it is possible that the populations of the Andes and, in particular, northern Colombia, are most threatened given the high rates of deforestation that affect these regions (
In Colombia, there is little information about the ecosystems occupied by V. spectrum. Herein, we recognize it in diverse ecosystems, but mainly associated with primary and secondary forests (
Some recent records confirm the low detectability and local rarity of the Spectral Bat. The new record for the Department of Antioquia confirms the presence of the species that has not been registered since 1959 for the Urabá subregion (
Forest cover is the most important factor behind V. spectrum’s distribution, and the response curves for this variable support this view – as has been suggested in other countries (
Although the ENM models produced high TSS and AUC scores, the ensemble maps should not be considered infallible. Factors such as human-wildlife conflict, competing species, dispersal limitations or stochastic processes can render climatically suitable areas unviable for V. spectrum. ENM methods can also be affected by unknown prevalence (the proportion of suitable sites occupied by the species), which can lead to misleading results (
Reviewed specimens show similar morphology and morphometrics, indicating little variability among the Colombian populations. The differences observed in the ear length and mean value of the forearm could be explained by the way the collectors historically took this measurement in the former and, in the latter, with clinal variation also observed in other bat species (
The Spectral Bat is listed as Near Threatened by IUCN (
The key localities with capacity to contribute to the long-term survival of this species are mostly outside the boundaries of Protected Areas. In particular, we highlight the lack of PAs in the Andean region, because it is the region with the largest area of distribution in Colombia and an area quite threatened by the increasing human land use and habitat transformation. Protected areas are considered a central axis of national and international conservation strategies, becoming a fundamental tool to ensure the conservation of species and their ecosystems (
Future studies should consider contributing to the knowledge of the natural history of this rare species. Widely unknown aspects such as habitat use and preference, population density, occupancy area and response to climate change should be a priority. The conservation of the Spectral Bat will depend on actions dedicated to the preservation and recovery of its habitat, to the formulation of Protected Areas and the development of environmental education activities. Likewise, the species should be included in plans of public or private areas where their presence has been recorded, as well as the protection of its identified shelters.
We thank the curators and assistants of the collections for providing data, and particularly the Kurupira Foundation for logistical and financial support. We extend our appreciation to Camilo Ramos Madera for his support in the revision of the document and Cristian Moreno for the distributional records map. Finally, we thank the reviewers for their suggestions and revision to the manuscript. Tatiana Toro-Sánchez by allowing the use of the photographs from Department of Tolima.
Natural History Museums examined. Reviewed personally * and online ** collections.
Colección de Mamíferos del Instituto Alexander von Humboldt (
Colección de Mamíferos de la Universidad de Nariño (
Colección Mastozoológica del Chocó (CMCH), Quibdó**
Colección de Mastozoología Universidad del Quindío (CMUQ), Armenia**
Colección Teriológica de la Universidad de Antioquia (
Colección Mastozoológica Universidad Industrial de Santander (UIS-MHN), Bucaramanga**
Colección Zoológica Universidad de Córdoba (CZUC), Montería*
Colección Zoológica Universidad del Tolima (CZUT), Ibagué*
Instituto de Ciencias Naturales (
Museo de Historia Natural Universidad Distrital Francisco José de Caldas (MUD), Bogotá*
Museo Javeriano de Historia Natural “Lorenzo Uribe, S.J.” (MPUJ), Bogotá*
Museo de La Salle (
Museo de Historia Natural Universidad de Los Andes (Andes-M), Bogotá*
Museo de Historia Natural Universidad de Caldas (MHN-UCa), Manizales*
Museo de Historia Natural Universidad del Cauca (
Universidad del Valle-Vertebrados Mamíferos (