Therya_notes_FE_9

THERYA NOTES 2022, Vol. 3 : 98-103 DOI: 10.12933/therya_notes-22-78

Documentation of a road-killed spectral bat (Vampyrum spectrum) and first report of the species in Tabasco, México

Documentación de un murciélago espectral (Vampyrum spectrum) atropellado en carretera y primer reporte de la especie en Tabasco, México

Coral Jazvel Pacheco-Figueroa1, Juan de Dios Valdez-Leal1*, Fernando A. Cervantes2, Ruth del Carmen Luna-Ruíz3, Ena Edith Mata-Zayas1,

and Joel C. Saenz4

1División Académica de Ciencias Biológicas de la UJAT. Carretera Villahermosa-Cárdenas Km 0.5, C. P. 86150. Villahermosa, Tabasco, México. E-mail: coral.pacheco@ujat.mx (CJP-F); juan.valdez@ujat.mx (JDV-L); ena.mata@ujat.mx (EEM-Z).

2Colección Nacional de Mamíferos, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México. Ciudad Universitaria, C. P. 04510. Ciudad de México. E-mail: fac@ib.unam.mx (FAC).

3Universidad Intercultural del Estado de Tabasco. Carretera Oxolotan Km 1 s/n, C. P. 86890. Tacotalpa, Tabasco, México. E-mail: bio.ruthclr@gmail.com (RCL-R).

4Instituto Internacional en Conservación y Manejo de Vida Silvestre (ICOMVIS), Universidad Nacional. Apartado 1350-3000. Heredia, Costa Rica. E-mail: jsaenz@una.ac.cr (JCS).

*Corresponding author

Roads are human infrastructures with adverse effects on wildlife; in the case of mammals, bats are rarely mentioned as road-killed animals. This work describes the record of a road-killed spectral bat (Vampyrum spectrum) on the El Bellote–Santa Cruz section of the 180 highway in Paraiso, Tabasco, México, the first report of the species in the state. The spectral bat is the largest bat species in the Americas, considered locally rare. The finding was recorded in 2015 during a wildlife mortality sampling tour. The specimen was collected and deposited in the National Mammals Collection of Universidad Nacional Autónoma de México. The corpse of a bird (Tyrannidae) was also found during the sampling trip. This is the first documented record of a road-killed spectral bat. The unique flight characteristics of the spectral bat, such as flying near obstacles and at ground level, make it vulnerable to vehicles and collisions. Mitigation measures should be applied to reduce the risk of collision for this species in particular and all bats in general.

Key words: Bats; mammals; road ecology; roadkill.

Las carreteras son infraestructuras humanas con efectos negativos para la fauna silvestre; en el caso de los mamíferos, los murciélagos pocas veces se mencionan como víctimas de atropellos. En este trabajo, se presenta el registro de un murciélago espectral (Vampyrum spectrum) atropellado en la carretera 180 del Tramo El Bellote-Santa Cruz, Paraíso, Tabasco, México, representando el primer reporte de esta especie en el estado. El murciélago espectral es el más grande del Continente Americano, el cual es considerado como una especie localmente rara. El hallazgo ocurrió en 2015 durante un recorrido de un muestreo de mortandad de fauna silvestre. El ejemplar fue recolectado e incorporado a la Colección Nacional de Mamíferos de la Universidad Nacional Autónoma de México. Al momento de la colecta se encontró también el cuerpo de un ave (Tyrannidae). El presente registro corresponde al primero de atropellamiento de murciélago espectral del que se tenga documentación. Las características particulares de vuelo del murciélago espectral de volar cerca de obstáculos y a ras del suelo, lo hace vulnerable a los vehículos y a las colisiones. Es necesario tomar medidas de mitigación que disminuyan los riesgos de colisión para esta especie y murciélagos en general.

Palabras clave: Atropello; ecología de carreteras; mamíferos; murciélagos.

Roads are the type of anthropic infrastructure with the most adverse effects on wildlife, mainly from collisions with vehicles and the impact of roads on habitat loss, fragmentation, and alteration (van der Ree et al. 2011; van der Ree et al. 2015; Schwartz et al. 2020; Benítez et al. 2021). Roadkills account for 6.25 % of vertebrate mortality (Hill et al. 2019), with large mammals as the most recorded and vulnerable taxa, followed by birds, amphibians, and reptiles (Rytwinski and Fahrig 2012; Silva et al. 2021).

Mammals are a group of great concern because of their vulnerability to local extinction due to roadkills (Grilo et al. 2021) and the human and economic costs of these accidents (Ascensão et al. 2021). The impact of roads on wild mammals has been focused mainly on ungulates, large carnivores, and small non-flying mammals (Novaes et al. 2018; Pinto et al. 2020).

Bats are rarely detected in roadkill studies compared to birds and amphibians (Berthinussen and Altringham 2012; Medinas et al. 2019; de Figueiredo Ramalho et al. 2021a; de Figueiredo Ramalho et al. 2021b). This group of mammals is vulnerable to roadkills due to the barrier effect limiting their displacement from one site to another. Despite its low detectability and scarce documentation, they are assumed to be at high risk of collision with motor vehicles given their flight habits by preferring to forage in the linear elements of the landscape (Bennett and Zurcher 2013; Claireau et al. 2021). They go unnoticed due to their small size, their ability to hide out among the vegetation, rapid decomposition of the corpse, and predation by scavengers (Russell et al. 2009; Teixeira et al. 2013; Altringham and Kerth 2016; Delgado et al. 2019). Since bats are less frequently detected than most vertebrates in road studies, collisions with vehicles might be an important cause of bat mortality (Lesiński et al. 2011).

The spectral bat (Vampyrum spectrum) is the largest bat species in the Americas, with a wingspan from 101 to 110 cm and a weight of approximately 180 gr (Navarro and Wilson 1982; Gardner 2008). It belongs to the family Phyllostomidae and is an apex predator with a wide home range and small scattered populations (Timm et al. 1989; Ceballos and Oliva 2005). This species inhabits caves or tree cavities and lives as solitary individuals or in family groups of about 5 members (Solari et al. 2019).

The spectral bat has a wide distribution in the Neotropical region, from Veracruz in México to Brazil, Perú, and Bolivia (Hall 1981; Medellín et al. 2008; Solari et al. 2019). This species has been found in tropical forests and secondary and riparian vegetation (Vehrencamp et al. 1977; Solari et al. 2019), and recorded in elevations from 4 m to 1,600 m (Timm et al. 1989; LaVal and Rodríguez 2002).

Vampyrum spectrum is a carnivorous bat (Gual-Suárez and Medellín 2021) that consumes rodents, other bats, probably insects, and birds of 20 gr to 150 gr. Up to 18 bird species have been identified in their diet, mainly trogons (Trogonidae), columbids (Columbidae), motmots (Momotidae), cuckoos (Cuculidae), wrens (Troglodytidae), and icterids (Icteridae; Timm et al. 1989; Reid 2006; Reid et al. 2010). Vampyrum spectrum is considered locally rare (Reid 2006). In México, it is classified as endangered (SEMARNAT 2010), while the International Union for Conservation of Nature listed it as Near Threatened (IUCN 2021). The main threats to their populations are fragmentation, habitat loss (Solari 2018), and anthropic destruction of their shelters due to negative perceptions influenced by myths and ignorance (Gómez-Nísino 2006).

This note reports for the first time one specimen of V. spectrum inhabiting the Tabasco plain that was road-killed from a collision with a vehicle on a road in the Paraiso municipality.

This information was obtained during the Wildlife Mortality Project in the Tabasco Plain conducted by researchers of the Universidad Juárez Autónoma de Tabasco (UJAT). The objective of this project was to identify critical sites of wildlife collisions and to define mitigation measures in the Tabasco Plain.

The prevailing climate in the study area is warm humid, with annual precipitation ranging from 1,500 to 2,000 mm and annual mean temperatures above 26 °C, with maximum values between 30 °C and 34 °C in May and minimum between 20 °C and 22 °C in January (Hernández-Santana et al. 2008). Dominant winds flow from E to NE at an average speed of 4 to 6 m/s (Hernández-Santana et al. 2008). There are three clearly defined seasons in a year: dry, rainy, and winter rainy seasons (Moguel and Molina-Enríquez 2000). The dominant habitats are mangrove forests, wetlands, cultivated pastures, and water bodies (INEGI 2016).

The prevailing land use at the collision site is permanent rainfed agriculture (INEGI 2016). The study section is a paved state road consisting of two C-type lanes with an average crown width of 18 m. The speed limit is 80 km/h, and the mean traffic volume is 103 vehicles per day, of which 83.6 % are cars. The Annual Mean Daily Traffic (TDPA, in Spanish) was 3,489 vehicles in the year of the road-kill event (SCT 2016).

The corpse was photographed, collected, and brought to the Landscape Ecology and Global Change Laboratory of the Biological Sciences Division at UJAT for identification. Afterward, it was frozen and sent to the National Mammal Collection (CNMA, in Spanish) of the Institute of Biology at Universidad Nacional Autónoma de México (UNAM), where its identification was confirmed. The specimen was conventionally prepared (skin and skeleton) and listed with the corresponding coordinates. The following measurements were recorded in millimeters (mm): total length (LT), tail length (LC); rear right leg length (LPDT); and right ear length (LOD). Additionally, the total weight was reported.

The sites where this species has been documented in México were gathered from a survey in the literature, the Global Biodiversity Information Facility (GBIF 2021), and the NaturaLista platform (2021). The data obtained was visualized on a map of the region.

On 16 May 2015, at 6:00 hr, a male road-killed V. spectrum bat was observed during a monitoring trip (Figure 1). The animal was found lying on the El Bellote-Santa Cruz section of the 180 highway in Chiltepec village, Paraiso municipality, Tabasco. It was recorded during the rainy season (May to October) when precipitation exceeds 120 mm. The collision occurred on a stretch of the straight road surrounded by secondary vegetation associated with fruit trees: mango (Magnifera indica), banana (Musa paradisiaca), coconut (Cocos nucifera), and gliricidia (Gliricidia sepium).

The bat specimen was recorded under catalog data CNMA 47872, male; sampling locality 0.64 km W Aquiles Serdán, Paraíso municipality, Tabasco (18° 24’ 46.89” N, 93° 0’ 57.74” W), on the Chiltepec-Vicente Guerrero highway, at 1 m above sea level. The measurements of the specimen (mm) were: LT 144; LC 0; LPDT 32.0; LOD 42. total weight was 182.9 gr.

The bat specimen was collected with bird feathers in the mouth. The headless body of a bird of the family Tyrannidae was also observed (Figure 1). The head was not found near the site, so the bird was likely captured by the bat.

Only 13 previous records of V. spectrum were found (Figure 2): 3 in Veracruz (Goldman 1917; Navarro 1979; GBIF 2021), 2 in Oaxaca (Alfaro et al. 2005; Santos Moreno et al. 2010), 3 in Chiapas (Hernández-Mijangos et al. 2008; López et al. 1998; Peña-Cuéllar et al. 2012), 4 in Campeche (Hernández-Huerta et al. 2000; Escalona-Segura et al. 2002; NaturaLista 2021a, 2021b), and 1 in Quintana Roo (NaturaLista 2021c). Therefore, this is the first documented record of the species for Tabasco, which is also a roadkill report.

Although bats are a well-studied group in Tabasco (Sánchez and Romero 1995; García-Morales et al. 2014; Hidalgo-Mihart et al. 2016), the presence of V. spectrum had not been previously reported in the literature. According to Solari et al. (2019), its distribution in México comprises southern Veracruz, Oaxaca, Chiapas, and the southern Yucatán Península, so this record adds Tabasco to its distribution range.

Bats are among the rarest organisms reported in roadkill studies (de Figueiredo Ramalho and Aguiar 2020; de Figueiredo Ramalho et al. 2021a; Vargas-Contreras et al. 2021), but an underestimation is plausible due to the poor detectability of these individuals. Furthermore, specific studies have reported that roads cause significant adverse effects on this group, particularly on insectivorous bats (Berthinussen and Altringham 2012; Medinas et al. 2019). The families most commonly recorded are Vespertilionidae and Phyllostomidae (de Figueiredo Ramalho and Aguiar 2020; Novaes et al. 2018).

In México, bat records in road ecology studies are scarce (González-Gallina and Badillo 2013; González-Gallina et al. 2013; Pacheco-Figueroa et al. 2021; Vargas-Contreras et al. 2021), probably because of the sampling techniques applied. As mentioned by Vargas-Contreras et al. (2021), corpses lying on the road are difficult to spot since the speed of the vehicle forces bats out of the road area and into the vegetation. Therefore, this is likely the first record of a road-killed V. spectrum (González-Gallina and Badillo 2013; González-Gallina et al. 2013; Ceron et al. 2017; Cervantes-Huerta et al. 2017; de Figueiredo Ramalho and Aguiar 2020; de Figueiredo Ramalho et al. 2021a).

Furthermore, the flight strategy influences the risk of collision (Vargas-Contreras et al. 2021). Species flying at less than 10 m high, such as members of the family Phyllostomidae, face a higher risk of vehicle collisions (de Figueiredo Ramalho and Aguiar 2020); this family includes the spectral bat, the largest in the Neotropics. The spectral bat uses smell to locate its prey (Veherencamp et al. 1977) and flies slowly, near obstacles, and at ground level (Khan 2015), making it particularly vulnerable to vehicles and collisions. Spectral bats are likely to move across roads continuously, being frequently hit by vehicles, but road surveys are not efficient enough to record them.

The impact of roads on bats remains poorly documented and with unknown effects on species that are rarely observed. Most of the available research focuses on four-lane, high-traffic roads, and large or charismatic species. Low-to-moderate traffic roads are considered to have negligible impacts. However, small roads run across landscapes all over the world, with poorly studied species incidentally found in collision events on these roads.

The spectral bat is threatened by habitat loss and the negative perception of bats among people (Ceballos and Oliva 2005; Gómes-Nísimo 2006; Solari 2018). This species prefers little-disturbed habitats; however, as Vleut (2013) mentioned, it can also take advantage of secondary vegetation, as long as there are forests that provide roost sites and shelter. Bats are most frequently hit by vehicles on roads close to vegetation that offers shelter, food, and roost sites (Rusell et al. 2009; Roemer et al. 2020). The road where the killed spectral bat was found is surrounded by natural vegetation dominated by mangrove trees, providing shelter and roost sites (Timm et al. 1989). Besides, the tyrannid bird, which apparently was caught as prey, suggests that these bats forage in patches adjacent to the road. The road studied represents high-risk conditions for the spectral bat to get hit by vehicles.

The road section where the bat was found showed a marked increase in the TDPA from 5,768 vehicles per year in 2015 to 6,977 vehicles per year in 2016. Although these values dropped in the following years (6,002 vehicles per year in 2019; SCT 2021), the risk of collision persists because this road is part of the Modernization and Expansion Plan of the Sánchez Magallanes – Paraíso – Frontera Highway (GET 2019), which involves expansion works and increased traffic flow, particularly cargo trucks.

Since V. spectrum is a threatened species, it should be considered in mitigation measures to reduce the impact of roads on this species. Without proper control and measures, isolated populations of V. spectrum in Tabasco may suffer adverse effects without even having an estimate of their populations.

Regarding mitigation measures, green bridges allow safe passage of bats (de Figueiredo Ramalho and Aguiar 2020), but proper measures must be applied to guide bats towards these structures. Underpasses such as large bridges and culverts are considered effective measures (Claireau et al. 2019); however, these passages are implemented mainly in temperate climates, so there is no evidence of their effectiveness in tropical areas (Vargas-Contreras et al. 2021). Therefore, these measures must be applied along with a monitoring program allowing for future adaptations.

The most effective measure is the reduction of speed limits in areas with tree cover surrounding the road because these are the sites with the highest probability of vehicle-wildlife collisions. Structural modifications should be implemented by planting trees or installing fences that force bats to fly higher (Vargas-Contreras et al. 2021). Continuous artificial light affects the behavior of nocturnal species (MITECO 20219); therefore, if the road infrastructure requires lighting, it should be kept to a minimum (Garin et al. 2016).

Acknowledgements

We thank the PFI Program of the UJAT for the funds granted for this study; to all the reviewers of the manuscript; to all the students and volunteers from the Wildlife Mortality Project in the Tabasco Plain for their collaboration. We also thank the assistance of the anonymous reviewers for their comments and suggestions. M. E. Sánchez-Salazar translated the manuscript into English.

Literature cited

Alfaro, A. M., et al. 2005. The false vampire bat Vampyrum spectrum in Oaxaca, México. Bat Research News 46:145-146.

Altringham, J., and G. Kerth. 2016. Bats and roads. Pp. 35-62 in Bats in the Anthropocene: conservation of bats in a changing world (Voigt, C. C., and T. Kingstons, eds.). Springer Cham Heidelberg. New York, Dordrecht, London.

Ascensão, F., et al. 2021. Preventing wildlife roadkill can offset mitigation investments in short-medium term. Biological Conservation 253:108902.

Benítez, A. A., et al. 2021. Tendencias del impacto de las vías de comunicación sobre la fauna silvestre: conclusiones para las ANP del sureste de México y previsiones ante el Tren Maya. Pp. 653-705 in Impacto de las vías de comunicación sobre la fauna silvestre en áreas protegidas: Estudios de caso para el sureste de México (Benítez, A. A., and G. Escalona-Segura, eds.). El Colegio de la Frontera Sur. Campeche, México.

Bennett, V. J., and A. A. Zurcher. 2013. When corridors collide: Road‐related disturbance in commuting bats. Journal of Wildlife Management 77:93-101.

Berthinussen, A., and J. Altringham. 2012. The effect of a major road on bat activity and diversity. Journal of Applied Ecology 49:82-89.

Ceballos, G., and G. Oliva. 2005. Los mamíferos silvestres de México. Fondo de Cultura Económica, Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. México City, México.

Ceron, K., et al. 2017. Roadkilled bats (Mammalia: Chiroptera) in two highways of Santa Catarina state, Southern Brazil. Oecologia Australis 21:207-212.

Cervantes-Huerta, R., et al. 2017. Atropellamiento de vertebrados en tres tipos de carreteras de la región montañosa central de Veracruz, México. Acta Zoológica Mexicana 33:472-481.

Claireau, F., et al. 2019. Major roads have important negative effects on insectivorous bat activity. Biological Conservation 235:53-62.

Claireau, F., et al. 2021. Bat overpasses as an alternative solution to restore habitat connectivity in the context of road requalification. Ecological Engineering 131:34-38.

de Figueiredo Ramalho, D., and L. Aguiar. 2020. Bats on the Road—A Review of the Impacts of Roads and Highways on Bats. Acta Chiropterologica 22:417-433.

de Figueiredo Ramalho, D., et al. 2021a. Factors influencing bat road casualties in a Neotropical savanna. Perspectives in Ecology and Conservation 19:189-194.

de Figueiredo Ramalho, et al. 2021b. Hit the road bat! High bat activity on the road verges in Brazilian savanna. Journal of Mammalogy 102:695-704.

Delgado, J. D., et al. 2019. The spatial distribution of animal casualties within a road corridor: Implications for roadkill monitoring in the southern Iberian rangelands. Transportation research part D: transport and environment 67:119-130.

Escalona-Segura, G., et al. 2002. Registros importantes de Mamíferos para Campeche, México. Revista Mexicana de Mastozoología (Nueva Época) 6:99-103.

García-Morales, R., et al. 2014. Las áreas naturales protegidas y su papel en la conservación de los murciélagos del estado de Tabasco, México. Therya 5:725-736.

Garin, I., et al. 2016. Guía para mitigar atropellos de Murciélagos. CEDR Programa Transnacional sobre Investigación Viaria Convocatoria 2013: Carreteras y Vida Silvestre. Conference of European Directors Roads. https://secemu.org/wp-content/uploads/2017/01/11-Guidelines_for_bat_mitigation_on_roads_Spanish.pdf. Accessed October 5, 2021.

Gardner, A. L. (ed.). 2008. Mammals of South America: Xenarthrans, Shrew and Bats (Vol. 1). The University of Chicago. Chicago, U.S.A.

GBIF (Global Biodiversity Information Facility). 2021. Occurrence Download. https://doi.org/10.15468/dl.husxv4.

GET (Gobierno del Estado de Tabasco). 2019. Plan Estatal de Desarrollo 2019-2024. Tabasco. https://tabasco.gob.mx/sites/default/files/users/planeacion_spf/PLED%202019-2024.pdf.

Goldman, E. A. 1917. New mammals from north and Middle America. Proceedings of the Biological Society of Washington 30:107-116.

Gómez-Nísino, A. 2006. Ficha técnica de Vampyrum spectrum. In: Medellín, R. (compilador). Los mamíferos mexicanos en riesgo de extinción según el PROY-NOM-059-ECOL-2000. Instituto de Ecología, Universidad Nacional Autónoma de México. Bases de datos SNIB-CONABIO. Proyecto No. W005. México City, México.

González, A., and G. B. Badillo. 2013. Road ecology studies for Mexico: A review. Oecologia Australis 17:175-190.

González-Gallina, A., et al. 2013. Los pequeños, los olvidados y los muertos: impacto de la carretera en los vertebrados y sus implicaciones para las estrategias de mitigación. Biodiversidad y Conservación 22:325-342.

Grilo, C., et al. 2021. Conservation threats from roadkill in the global road network. Global Ecology and Biogeography 30:2200-2210.

Gual-Suárez, F., and R. A. Medellín. 2021. We eat meat: A review of carnivory in bats. Mammal Review 51:540-558.

Hall, E. R. 1981. Mammals of North America. Wiley. New York, U.S.A.

Hernández-Huerta, A., et al. 2000. Records of small mammals in the Calakmul Biosphere Reserve, Yucatán Peninsula. The Southwestern Naturalist 45:340-344.

Hernández-Mijangos, L. A., et al. 2008. Nuevas localidades en la distribución de murciélagos filostóminos (Chiroptera: Phyllostomidae) en Chiapas, México. Revista Mexicana de Mastozoología (Nueva Época) 12:163-169.

Hernández-Santana, J. R., et al. 2008. Morfodinámica de la línea de costa del estado de Tabasco, México: tendencias desde la segunda mitad del siglo XX hasta el presente. Investigaciones Geográficas 65:7-21.

Hidalgo-Mihart, M. G., et al. 2016. Mamíferos del estado de Tabasco. Pp. 441-472 in Riqueza y Conservación de los Mamíferos en México a Nivel Estatal (Briones-Salas, M., Y. Hortelano-Moncada, G. Magaña-Cota, G. Sánchez-Rojas, and J. E. Sosa-Escalante, eds.). Instituto de Biología, Universidad Nacional Autónoma de México, Asociación Mexicana de Mastozoología A. C., Universidad de Guanajuato. México City, México.

Hill, J. E., et al. 2019. Cause‐specific mortality of the world’s terrestrial vertebrates. Global Ecology and Biogeography 28:680-689.

Instituto Nacional de Estadística y Geografía (INEGI). 2016. 1:250,000. Conjunto de datos vectoriales de uso de suelo y vegetación serie VI. https://www.inegi.org.mx/app/biblioteca/ficha.html?upc=889463173359.

International Union for Conservation of Nature (IUCN). 2021. The IUCN Red List of Threatened Species. Version 2021-1. https://www.iucnredlist.org.

Khan, V. 2015. The Online Guide to the Animals of Trinidad and Tobago. Vampyrum spectrum (Spectral Bat). The University of the West Indies, St. Augustine. Trinidad y Tobago. https://sta.uwi.edu/fst/lifesciences/sites/default/files/lifesciences/documents/ogatt/Vampyrum_spectrum%20-%20Spectral%20Bat.pdf.

LaVal, R. K., and B. Rodríguez. 2002. Murciélagos de Costa Rica. Editorial INBio. Santo Domingo, Heredia, Costa Rica.

Lesiński, G., et al. 2011. Bat casualties on a road crossing a mosaic landscape. European Journal of Wildlife Research 57:217-223.

López, T. M. C., et al. 1998. Vampyrum spectrum en Chiapas, México. Revista Mexicana de Mastozoología 3:135-136.

Medellín, R. A., et al. 2008. Identificación de los murciélagos de México. Clave de campo. 2a Ed. Instituto de Ecología, Universidad Nacional Autónoma de México. México City, México.

Medinas, D., et al. 2019. Road effects on bat activity depend on surrounding habitat type. Science of The Total Environment 660:340-347.

Ministry for Ecological Transition and Demographic Challenge (MITECO). 2021. Edge and verge effects of transport infrastructure. Mitigating their Impact on Biodiversity. Documents for the Mitigation of Habitat Fragmentation Caused by Transport Infrastructure, 7. Ministry for Ecological Transition and Demographic Challenge. Madrid, Spain.

Moguel, O. E. J., and M. J. F. Molina-Enríquez. 2000. La precipitación pluvial en Tabasco y Chiapas. Kuxulkab 5:1-8.

Naturalista. 2021a. Download August 10, 2021. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. https://www.naturalista.mx/observations/38868037. Observación de Jaime Marcelo Aranda (Marcelo_Aranda), México.

Naturalista. 2021b. Download August 10, 2021. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. https://www.inaturalist.org/observations/4032127. Observación de Rodrigo Medellin (Medellin), México.

Naturalista. 2021c. Download August 10, 2021. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. https://www.naturalista.mx/observations/69890689. Observación de Erika Jasmin Cruz-Basan (erika_cruz), México.

Navarro, L. D. 1979. Vampyrum spectrum (Chiroptera, Phyllostomatidae) in Mexico. Journal of Mammalogy 60:435.

Navarro, D., and D. E. Wilson. 1982. Vampyrum spectrum. Mammalian Species 184:1-4.

Novaes, R. L. M., et al. 2018. On a collision course: the vulnerability of bats to roadkills in Brazil. Mastozoología Neotropical 25:115-128.

Pacheco-Figueroa, C. J., et al. 2021. Puntos de alta siniestralidad de vertebrados en la carretera costera de Tabasco. Pp. 348-367 in Impacto de las vías de comunicación sobre la fauna silvestre en áreas protegidas: Estudios de caso para el sureste de México (Benítez, A. A., and G. Escalona-Segura, eds.). El Colegio de la Frontera Sur. Campeche, México.

Peña-Cuéllar, E., et al. 2012. Phyllostomid bat assemblages in different successional stages of tropical rain forest in Chiapas, Mexico. Biodiversity and Conservation 21:1381-1397.

Pinto, F. A., et al. 2020. Effects of roads on terrestrial vertebrate species in Latin America. Environmental Impact Assessment Review 81:106337.

Reid, F. 2006. A field guide to mammals of North America, North of Mexico (Vol. 4). Houghton Mifflin Harcourt. New York, U.S.A.

Reid, F., et al. 2010. The wildlife of Costa Rica: A Field Guide. A Zona Tropical Publication. Cornell University Press. Ithaca, New York, U.S.A.

Roemer, C., et al. 2020. Influence of local landscape and time of year on bat-road collision risks. Peer Community Journal 1:e54.

Russell, A. L., et al. 2009. Road-killed bats, highway design, and the commuting ecology of bats. Endangered Species Research 8:49-60.

Rytwinski, T., and L. Fahrig. 2012. Do species life history traits explain population responses to roads? A meta-analysis. Biological Conservation 147:87-98.

Sánchez, H., C, and M. D. L. Romero A. 1995. Murciélagos de Tabasco y Campeche: una propuesta para su conservación (Vol. 24). Universidad Nacional Autónoma de México. México City, México.

Santos-Moreno, A., et al. 2010. Records of bats from Oaxaca, Mexico. The Southwestern Naturalist 55:454-456.

Schwartz, A. L., F. M. Shilling, and S. E. Perkins. 2020. The value of monitoring wildlife roadkill. European Journal of Wildlife Research 66:1-12.

Secretaría de Comunicaciones y Transportes (SCT). 2016. Datos Viales 2016. Subsecretaria de Infraestructura. México City, México.

Secretaría de Comunicaciones y Transportes (SCT). 2021. Datos Viales 2020. Volúmenes de Tránsito en la Red Nacional de Carreteras Pavimentadas. México. http://datosviales2020.routedev.mx/main#. Accessed August 1, 2021.

Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT). 2010. Norma Oficial Mexicana NOM-059-SEMARNAT-2010, Protección ambiental - Especies nativas de México de Flora y Fauna Silvestres-Categorías de Riesgo y especificaciones para su inclusión, exclusión o cambio – Lista de Especies en Riesgo. Diario Oficial de la Federación 2ª Sección, 30 de diciembre del 2010. MODIFICACIÓN del Anexo Normativo III, 19 de Noviembre del 2019.

Silva, I., et al. 2021. The road less traveled: addressing reproducibility and conservation priorities of wildlife-vehicle collision studies in tropical and subtropical regions. Global Ecology and Conservation 27:e01584.

Solari, S. 2018. Vampyrum spectrum. The IUCN Red List of Threatened Species 2018: e.T22843A22059426. https://dx.doi.org/10.2305/IUCN.UK.2018-2.RLTS.T22843A22059426.

Solari, S., et al. 2019. Family Phyllostomidae (New World Leaf-Nosed Bats). Pp. 444-583 in Handbook of the mammals of the world. Vol. 9. Bats (Wilson, D. E., and Mitermmeier, R. A., eds.). Lynx Edicions. Barcelona, Spain.

Teixeira, F. Z., et al. 2013. Vertebrate Road mortality estimates: effects of sampling methods and carcass removal. Biological Conservation 157:317-323.

Timm, R. M., et al. 1989. Mammals of the La Selva – Braulio Carrillo Complex, Costa Rica. North American Fauna. 75. Washington, D. C., U.S.A.

van der Ree, R., et al. 2011. Effects of roads and traffic on wildlife populations and landscape function: road ecology is moving toward larger scales. Ecology and Society 16:48.

van der Ree, R., et al. 2015. The ecological effects of linear infrastructure and traffic: challenges and opportunities of rapid global growth. Pp. 1-9 in Handbook of Road Ecology (Redd, R., D. J. Smith, and C. Grilo, eds.). John Wiley and Sons Ltd. West Sessex, United Kingdom.

Vargas-Contreras, J. A, et al. 2021. Impacto por atropellamiento cerca de la cueva el Volcán de los Murciélagos, en Calakmul, Campeche, México. Pp. 320-347 in Impacto de las vías de comunicación sobre la fauna silvestre en áreas protegidas: Estudios de caso para el sureste de México (Benítez, A. A., and G. Escalona-Segura, eds.). El Colegio de la Frontera Sur. Campeche, México.

Vehrencamp, S. L, et al. 1977. Observations on the foraging behavior and avian prey of the Neotropical carnivorous bat, Vampyrum spectrum. Journal of Mammalogy 58:469-478.

Vleut, I. J. 2013. Factores determinantes sobre la presencia de murciélagos en bosques secundarios bajo manejo tradicional en Lacanhá, Selva Lacandona. Chiapas, México. Tesis Doctoral. El Colegio de la Frontera Sur. San Cristóbal de Las Casas, Chiapas, México.

Associated editor: Cristian Kraker-Castañeda

Submitted: October 5, 2021; Reviewed: April 19, 2022.

Accepted: April 28, 2022; Published on line: May 5, 2022.