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Vampire bat

(Redirected fromVampire bats)
For the 1933 film, seeThe Vampire Bat.For the 2005 television film, seeVampire Bats (film).

Vampire bats,members of the subfamilyDesmodontinae,areleaf-nosed batscurrently found in Central and South America. Their food source is the blood of other animals, a dietary trait calledhematophagy.Three extant batspeciesfeed solely on blood: thecommon vampire bat(Desmodus rotundus), thehairy-legged vampire bat(Diphylla ecaudata), and thewhite-winged vampire bat(Diaemus youngi). Two extinct species of the genusDesmodushave been found inNorth America.

Vampire bat
The image depicts the common vampire bat (i.e. Desmodus rotundus) hanging from a cave wall and staring at the camera.
Common vampire bat(Desmodus rotundus)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Chiroptera
Family: Phyllostomidae
Subfamily: Desmodontinae
Bonaparte,1845
Genera

Taxonomy

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Due to differences among the three species, each has been placed within a differentgenus,each consisting of one extant species. In the older literature, these three genera were placed within afamilyof their own, Desmodontidae, buttaxonomistshave now grouped them as asubfamily,Desmodontinae, in the New World leaf-nosed bat family,Phyllostomidae.[1]

The three known species of vampire bats all seem more similar to one another than to any other species. That suggests thathematophagyevolved only once, and the three species share this common ancestor.[1]: 163–167 

The placement of the three genera of the subfamily Desmodontinae within the New World leaf-nosed bat family Phyllostomidae Gray, 1825, may be summarized as:[2]

Evolution

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Vampire bats are ina diverse family of batsthat consume many food sources, includingnectar,pollen,insects,fruitandmeat.[1] The three species of vampire bats are the only mammals that have evolved to feed exclusively on blood (hematophagy) asmicropredators,a strategy withinparasitism.[4][5] Hematophagy is uncommon due to the number of challenges to overcome for success: a large volume of liquid potentially overwhelming thekidneys and bladder,[6]the risk ofiron poisoning,[7]and coping withexcess protein.[8] There are multiple hypotheses for how vampire bats evolved.

  • They evolved from frugivorous bats with sharp teeth specialized for piercing fruit[9]
  • They initially fed on theectoparasitesof large mammals, and then progressed to feeding on the mammals themselves[10](similar tored-billed oxpeckerfeeding behavior)
  • They initially fed on insects that were attracted to the wounds of animals, and then progressed to feeding on the wounds[11]
  • They initially preyed on small arboreal vertebrates[12]
  • They were arboreal omnivores themselves and began ingesting blood and flesh from wound sites of larger animals[13]
  • They were specialized nectar-feeders that evolved to feed on another type of liquid[14]

The vampire bat lineage diverged from its family 26 million years ago.[15]Thehairy-legged vampire batlikely diverged from the other two species of vampire bats 21.7 million years ago.[15] Because the hairy-legged vampire bat feeds on bird blood and it is themost basalof living vampire bats, it is considered likely that the first vampire bats fed on bird blood as well.[15] Recent analyses suggest that vampire bats arose from insectivores, which discount the frugivore, carnivore, and nectarivore hypotheses of origin.[15]Within 4 million years of diverging from other Phyllostomidae, vampire bats had evolved all necessary adaptations for blood-feeding, making it one of the fastest examples of natural selection among mammals.[15]

Anatomy and physiology

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A vampire bat skeleton, showing the distinctive incisors and canines

Unlike fruit bats, the vampire bats have short, conical muzzles. They also lack a nose leaf, instead having naked pads with U-shaped grooves at the tip. The common vampire bat,Desmodus rotundus,also has specializedthermoreceptorson its nose,[16]which aid the animal in locating areas where the blood flows close to the skin of its prey. A nucleus has been found in the brain of vampire bats that has a similar position and similarhistologyto theinfrared receptorof infrared-sensingsnakes.[17][18]

A vampire bat has front teeth that are specialized for cutting and back teeth that are much smaller than in other bats. Theinferior colliculus,the part of the bat's brain that processes sound, is well adapted to detecting the regular breathing sounds of sleeping animals that serve as its main food source.[19][20]

While other bats have almost lost the ability to maneuver on land, vampire bats can walk, jump, and even run by using a unique, bounding gait, in which the forelimbs instead of the hindlimbs are recruited for force production, as the wings are much more powerful than the legs. This ability to run seems to have evolved independently within the bat lineage.[21]

Vampire bats also have a high level of resistance to a group of bloodborne viruses known asendogenous retroviruses,which insert copies of their genetic material into their host's genome.[22]

It was recently discovered that the vampire bat's loss of the REP15 gene allows for enhanced iron secretion in adaptation to the high iron diet.[23]

Vampire bats use infrared radiation to locate blood hotspots on their prey. A recent study has shown that common vampire bats tune aTRP-channelthat is already heat-sensitive, TRPV1, by lowering its thermal activation threshold to about 30 °C (86 °F). This is achieved through alternative splicing of TRPV1 transcripts to produce a channel with a truncated carboxy-terminal cytoplasmic domain. These splicing events occur exclusively in trigeminal ganglia, and not in dorsal root ganglia, thereby maintaining a role for TRPV1 as a detector of noxious heat in somatic afferents.[24]The only other knownvertebratescapable of detectinginfrared radiationareboas,Python sandpit vipers,all of which have pit organs.

Ecology and life cycle

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Vampire bats tend to live in colonies in almost completely dark places, such as caves, old wells, hollow trees, and buildings. They range in Central to South America and live in arid to humid, tropical and subtropical areas. Vampire bat colony numbers can range from single digits to hundreds in roosting sites. The basic social structure of roosting bats is made of female groups and their offspring, a few adult males, known as "resident males", and a separate group of males, known as "nonresident males".[25]In hairy-legged vampire bats, the hierarchical segregation of nonresident males appears less strict than in common vampire bats.[26]Nonresident males are accepted into the harems when the ambient temperature lowers. This behavior suggests social thermoregulation.

Resident males mate with the females in their harems, and it is less common for outside males to copulate with the females.[25]Female offspring often remain in their natal groups.[25]Severalmatrilinescan be found in a group, as unrelated females regularly join groups.[25]Male offspring tend to live in their natal groups until they are about two years old, sometimes being forcibly expelled by the resident adult males.[25]Vampire bats on average live about nine years when they are in their natural environment in the wild.[27]

Vampire bats form strong bonds with other members of the colony. A related unique adaptation of vampire bats is the sharing of food. A vampire bat can only survive about two days without feeding, yet they cannot be guaranteed of finding food every night. This poses a problem, so when a bat fails to find food, it will often "beg" another bat for food. A "donor" bat may regurgitate a small amount of blood to sustain the other member of the colony. For equally familiar bats, the predictive capacity of reciprocity surpasses that of relatedness.[28]This finding suggests that vampire bats are capable of preferentially aiding their relatives, but that they may benefit more from forming reciprocal, cooperative relationships with relatives and non-relatives alike.[28]Furthermore, donor bats were more likely to approach starving bats and initiate the food sharing. When individuals of a population are lost, bats with a larger number of mutual donors tend to offset their own energetic costs at a higher rate than bats that fed less of the colony before the removal. Individuals that spend their own energy as a social investment of sorts are more likely to thrive, and higher rates of survival incentivize the behavior and reinforce the importance of large social networks in colonies.[29]These findings contradict the harassment hypothesis—which claims that individuals share food in order to limit harassment by begging individuals.[28]All considered, vampire bat research should be interpreted cautiously as much of the evidence is correlational and still requires further testing.[30]

Another ability that some vampire bats possess is identifying and monitoring the positions of conspecifics (individuals of the same species) simply byantiphonalcalling.[31]Similar in nature to the sound mother bats make to call to their pups, these calls tend to vary on a bat to bat basis which may help other bats identify individuals both in and outside of their roost.[32]

Vampire bats also engage insocial grooming.[33]It usually occurs between females and their offspring, but it is also significant between adult females. Social grooming is mostly associated with food sharing.[33]

Feeding

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A vampire bat feeding on a pig (taxidermy specimens)

Vampire bats hunt only when it is fully dark. Like fruit-eating bats, and unlike insectivorous and fish-eating bats, they emit only low-energy sound pulses. The common vampire bat feeds primarily on the blood of mammals (occasionally including humans), whereas both the hairy-legged vampire bat and white-winged vampire bat feed primarily on the blood of birds. Once the common vampire bat locates a host, such as a sleeping mammal, it lands and approaches it on the ground while on all fours. It then likely uses thermoception to identify a warm spot on the skin to bite. They then start to lick the area over and over again to make the place tender so it's easier to bite. Then create a small incision with their teeth and lap up blood from the wound.

Vampire bats, like snakes, have developed highly sensitive thermosensation, with specialized systems for detecting infrared radiation. Snakes co-opt a non-heat-sensitive channel, vertebrateTRPA1(transient receptor potential cation channel A1), to produce an infrared detector. However, vampire bats tune a channel that is already heat-sensitive,TRPV1,by lowering its thermal activation threshold to about 30 °C (86 °F), which allows them to sense the target.[34]

As noted by Arthur M. Greenhall:

The most common species, the common vampire (Desmodus) is not fastidious and will attack any warm-blooded animal. The white-winged vampire (Diaemus) appears to have a special preference for birds and goats. In the laboratory it has not been possible to feedDiaemuson cattle blood.[35]

If there is fur on the skin of the host, the common vampire bat uses itscanineandcheek teethlike a barber's blades to shave away the hairs. The bat's razor-sharpupper incisor teeththen make a 7 mm wide and 8 mm deep cut. The upper incisors lackenamel,which keeps them permanently razor sharp.[36]Their teeth are so sharp, even handling their skulls in a museum can result in cuts.[37]

The bat'ssaliva,left in the victim's resulting bite wound, has a key function in feeding from the wound. The saliva contains several compounds that prolong bleeding, such asanticoagulantsthat inhibit blood clotting,[38]and compounds that prevent the constriction of blood vessels near the wound.

Digestion

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A typical female vampire bat weighs 40 grams (1.4 oz) and can consume over 20 grams (1 fluid ounce) of blood in a 20-minute feed. This feeding behavior is facilitated by itsanatomyand physiology for rapid processing and digestion of the blood to enable the animal to take flight soon after the feeding. The stomach and intestine rapidly absorb the water in the blood meal, which is quickly transported to thekidneys,and on to thebladderfor excretion.[39][40]A common vampire bat begins to expel urine within two minutes of feeding. While shedding much of the blood's liquid facilitates flight takeoff, the bat still has added almost 20–30% of its body weight in blood. To take off from the ground, the bat generates extra lift by crouching and flinging itself into the air.[41]Typically, within two hours of setting out in search of food, the common vampire bat returns to its roost and settles down to spend the rest of the night digesting its meal. Digestion is aided by theirmicrobiome,and their genome protects them against pathogens in the blood.[42]Its stool is roughly the same as that from bats eating fruits or insects.[43]

Metabolism

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In a 2024 study published inBiology Letters,researchers explored how vampire bats generate energy from their blood meals, hypothesizing that they metabolizeamino acidsdue to their low-carbohydrate and low-fat diet. The team captured two dozen vampire bats inBelizeand fed them cow blood enriched withglycineandleucine.After consumption, the bats were placed on a treadmill for up to 90 minutes, during which breath samples were collected to measure oxygen intake andcarbon dioxideoutput. The results revealed that up to 60% of the bats’ energy production during exercise came from the rapid breakdown of these amino acids, revealing their ability to convert proteins into usable energy within ten minutes. Michael Hiller, a researcher at the LOEWE Center for Translational Biodiversity Genomics in Frankfurt, noted that this rapid metabolization of amino acids is "unparalleled in mammals" and described it as a compelling example ofconvergent evolution,where both vampire bats and blood-feeding insects developed similar strategies to adapt to their extreme diets.[44]

This metabolic specialization presents drawbacks, as vampire bats have diminished their ability to store alternative energy sources, rendering them susceptible tostarvationif they experience prolonged periods without feeding. To counteract this vulnerability, vampire bats engage inreciprocal altruism,regurgitating blood to assist conspecifics in need.[44]

Human health

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Common vampire batat theLouisville Zoo

Rabies

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Rabiescan be transmitted to humans and other animals by vampire bat bites. Since dogs are now widelyimmunized against rabies,the number of human rabies transmissions by vampire bats exceeds those by dogs in Latin America, with 55 documented cases in 2005.[45]The risk of infection to the human population is less than to livestock exposed to bat bites.[46]Various estimates of the prevalence of rabies in bat populations have been made; it has been estimated that less than 1% of wild bats in regions where rabies isendemicare infected with the virus at any given time.[47]Bats that are infected may be clumsy, disoriented, and unable to fly.[48]

Anticoagulant drug

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The unique properties of vampire bat saliva have found some positive use in medicine.

Various studies published inStroke: Journal of the American Heart Associationon a genetically engineered drug calleddesmoteplasewhich uses theanticoagulantproperties of thesalivaofDesmodus rotundusfound that it increased blood flow instrokepatients.[49]

See also

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Footnotes

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  1. ^abcWetterer, Andrea L.; Rockman, Matthew V.; Simmons, Nancy B. (2000)."Phylogeny of phyllostomid bats (Mammalia: Chiroptera): data from diverse morphological systems, sex chromosomes, and restriction sites"(PDF).Bull. Am. Mus. Nat. Hist.248:1–200.doi:10.1206/0003-0090(2000)248<0001:popbmc>2.0.co;2.hdl:2246/1595.S2CID83617355.Archived fromthe original(PDF)on 2014-02-03.Retrieved2013-02-22.
  2. ^Simmons, N.B.(2005)."Order Chiroptera".InWilson, D.E.;Reeder, D.M (eds.).Mammal Species of the World: A Taxonomic and Geographic Reference(3rd ed.). Johns Hopkins University Press. pp. 312–529.ISBN978-0-8018-8221-0.OCLC62265494.
  3. ^"Fossilworks: Desmodus".fossilworks.org.Retrieved17 December2021.
  4. ^Botero-Castro, Fidel; Tilak, Marie-Ka; Justy, Fabienne; Catzeflis, Francois; Delsuc, Frédéric; Douzery, Emmanuel J.P. (2018)."In cold blood: Compositional Bias and Positive Selection Drive the High Evolutionary Rate of Vampire Bats Mitochondrial Genomes".Genome Biology and Evolution.10(9): 2218–2239.doi:10.1093/gbe/evy120.PMC6127110.PMID29931241.
  5. ^Poulin, Robert;Randhawa, Haseeb S. (February 2015)."Evolution of parasitism along convergent lines: from ecology to genomics".Parasitology.142(Suppl 1): S6–S15.doi:10.1017/S0031182013001674.PMC4413784.PMID24229807.
  6. ^Breidenstein C. P. (1982). "Digestion and assimilation of bovine blood by a vampire bat (Desmodus rotundus) ".Journal of Mammalogy.63(3): 482–484.doi:10.2307/1380446.JSTOR1380446.
  7. ^Morton D.; Wimsatt W. A. (1980). "Distribution of iron in the gastrointestinal tract of the common vampire bat: Evidence for macrophage‐linked iron clearance".The Anatomical Record.198(2): 183–192.doi:10.1002/ar.1091980206.PMID7212303.S2CID13463459.
  8. ^Singer M. A. (2002). "Vampire bat, shrew, and bear: comparative physiology and chronic renal failure".American Journal of Physiology. Regulatory, Integrative and Comparative Physiology.282(6): R1583–R1592.doi:10.1152/ajpregu.00711.2001.PMID12010738.
  9. ^Slaughter, B. H. (1970). "Evolutionary trends of chiropteran dentitions".About Bats.Dallas: Southern Methodist University Press. pp. 51–83.
  10. ^Gillette, D. D. (1975). "Evolution of feeding strategies in bats".Tebiwa.18:39–48.
  11. ^Fenton M. B. (1992). "Wounds and the origin of blood‐feeding in bats".Biological Journal of the Linnean Society.47(2): 161–171.doi:10.1111/j.1095-8312.1992.tb00662.x.
  12. ^Sazima I (1978). "Vertebrates as food items of the woolly false vampire,Chrotopterus auritus".Journal of Mammalogy.59(3): 617–618.doi:10.2307/1380238.JSTOR1380238.
  13. ^Schutt, W. A., Jr. (1998). "Chiropteran hindlimb morphology and the origin of blood-feeding in bats". In T. H. Kunz, and P. A. Racey (eds.),Bat biology and conservation.Washington D.C.: Smithsonian Inst. pp. 157–168.ISBN978-1560988250
  14. ^Baker, Robert J.; Carter, Dilford C.; Jones, J. Knox. (1976).Biology of bats of the New World family Phyllostomatidae /.doi:10.5962/bhl.title.142603.
  15. ^abcdeBaker, R. J.; Bininda-Emonds, O. R.; Mantilla-Meluk, H.; Porter, C. A.; Van Den Bussche, R. A. (2012). "Molecular timescale of diversification of feeding strategy and morphology in New World leaf-nosed bats (Phyllostomidae): a phylogenetic perspective". In Gunnell, Gregg F; Simmons, Nancy B (eds.).Evolutionary history of bats: fossils, molecules and morphology.pp. 385–409.doi:10.1017/CBO9781139045599.012.ISBN9781139045599.
  16. ^Kürten, Ludwig; Schmidt, Uwe; Schäfer, Klaus (1984). "Warm and Cold Receptors in the Nose of the Vampire BatDesmodus rotundus".Naturwissenschaften.71(6): 327–328.Bibcode:1984NW.....71..327K.doi:10.1007/BF00396621.PMID6472483.S2CID31899356.
  17. ^Campbell, Angela L.; Naik, Rajesh R.; Sowards, Laura; Stone, Morley O. (2002)."Biological infrared imaging and sensing"(PDF).Micron.33(2): 211–225.doi:10.1016/S0968-4328(01)00010-5.PMID11567889.Archived fromthe original(PDF)on 2003-06-15.
  18. ^Kishida R, Goris RC, Terashima S, Dubbeldam J (1984). "A suspected infrared-recipient nucleus in the brainstem of the vampire bat,Desmodus rotundus".Brain Res.322(2): 351–355.doi:10.1016/0006-8993(84)90132-X.PMID6509324.S2CID33339192.
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  20. ^Gröger, Udo & Wiegrebe, Lutz (2006)."Classification of human breathing sounds by the common vampire bat,Desmodus rotundus".BMC Biology.4:18.doi:10.1186/1741-7007-4-18.PMC1513255.PMID16780579.
  21. ^Riskin, Daniel K.; Hermanson, John W. (2005)."Independent evolution of running in vampire bats".Nature.434(7031): 292.doi:10.1038/434292a.PMID15772640.S2CID4406312.
  22. ^Arnold, Carrie (22 February 2018)."Vampire Bats Survive by Only Eating Blood—Now We Know How".National Geographic.Archived fromthe originalon 18 September 2021.
  23. ^Moritz Blumer et al.,Gene losses in the common vampire bat illuminate molecular adaptations to blood feeding.Sci. Adv.8,eabm6494(2022).DOI:10.1126/sciadv.abm6494
  24. ^Gracheva, Elena O.; Cordero-Morales, Julio F.; González-Carcacía, José A.; Ingolia, Nicholas T.; Manno, Carlo; Aranguren, Carla I.; Weissman, Jonathan S.; Julius, David (2011)."Ganglion-specific splicing of TRPV1 underlies infrared sensation in vampire bats".Nature.476(7358): 88–91.doi:10.1038/nature10245.PMC3535012.PMID21814281.
  25. ^abcdeWilkinson G. S. (1985)."The Social Organization of the Common Vampire Bat II:Mating System, Genetic Structure and Relatedness".Behavioral Ecology and Sociobiology.17(2): 123–134.doi:10.1007/BF00299244.JSTOR4599815.S2CID12460893.
  26. ^Delpietro H. A., Russo R. G. (2002). "Observations of the common vampire bat (Desmodus rotundus) and the hairy-legged vampire bat (Diphylla ecaudata) in captivity ".Mammalian Biology – Zeitschrift für Säugetierkunde.67(2): 65–78.doi:10.1078/1616-5047-00011.
  27. ^"Vampire Bat".Animals.2014-03-01.Retrieved2022-12-05.
  28. ^abcCarter, G. G. & Wilkinson, G. S. (2013)."Food sharing in vampire bats: reciprocal help predicts donations more than relatedness or harassment".Proc R Soc B.280(1753): 20122573.doi:10.1098/rspb.2012.2573.PMC3574350.PMID23282995.
  29. ^Carter, Gerald; Farine, Damien; Wilkinson, Gerald (2017-05-01)."Social bet-hedging in vampire bats".Biology Letters.13(5): 20170112.doi:10.1098/rsbl.2017.0112.PMC5454239.PMID28539459.
  30. ^Carter, G. & Wilkinson, G. (2013)."Does food sharing in vampire bats demonstrate reciprocity?".Communicative and Integrative Biology.6(6): e25783.doi:10.4161/cib.25783.PMC3913674.PMID24505498.
  31. ^Carter, G. G.; Fenton, M. B.; Faure, P. A. (2009). "White-winged vampire bats (Diaemus youngi) exchange contact calls ".Canadian Journal of Zoology.87(7): 604–608.doi:10.1139/Z09-051.
  32. ^Carter, Gerald G.; Skowronski, Mark D.; Faure, Paul A.; Fenton, Brock (2008)."Antiphonal calling allows individual discrimination in white-winged vampire bats".Animal Behaviour.76(4): 1343–1355.doi:10.1016/j.anbehav.2008.04.023.S2CID53147832.Retrieved2018-12-01.
  33. ^abWilkinson G. S. (1986). "Social grooming in the common vampire bat,Desmodus rotundus".Animal Behaviour.34(6): 1880–1889.CiteSeerX10.1.1.539.5104.doi:10.1016/s0003-3472(86)80274-3.S2CID11214563.
  34. ^Gracheva, Elena (August 4, 2011)."Ganglion-specific splicing of TRPV1 underlies infrared sensation in vampire bats".Nature.476(7358): 88–91.doi:10.1038/nature10245.PMC3535012.PMID21814281.
  35. ^Greenhall, Arthur M. (1961).Bats in Agriculture,p. 8. A Ministry of Agriculture Publication. Trinidad and Tobago.
  36. ^Greenhall, Arthur M. (1988) "Feeding Behavior". In:Natural History of Vampire Bats(ed. by A. M. Greenhall and U. Schmidt), pp. 111–132. Boca Raton, FL: CRC Press.ISBN978-0-8493-6750-2
  37. ^Callaway, Ewen (October 31, 2008)."How vampires evolved to live on blood alone".New Scientist.Reed Business Information Ltd."You can actually cut yourself handling a bat skull in a museum, they're that sharp"
  38. ^Hawkey, Christine (1966). "Plasminogen Activator in Saliva of the Vampire BatDesmodus rotundus".Nature.211(5047): 434–435.Bibcode:1966Natur.211..434H.doi:10.1038/211434c0.PMID5967844.S2CID5301723.
  39. ^Price E. R.; Brun A.; Gontero-Fourcade M.; Fernández-Marinone G.; Cruz-Neto A. P.; Karasov W. H.; Caviedes-Vidal E. (2015). "Intestinal Water Absorption Varies with Expected Dietary Water Load among Bats but Does Not Drive Paracellular Nutrient Absorption".Physiol. Biochem. Zool.88(6): 680–684.doi:10.1086/683114.hdl:11336/14629.PMID26658415.S2CID206003403.
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  42. ^Katz, Brigit (23 February 2018)."How Vampire Bats Can Survive on a Diet of Blood".Smithsonian.Retrieved23 February2018.
  43. ^Emerson, Justin K.; Roark, Alison M. (April 2007). "Composition of guano produced by frugivorous, sanguivorous, and insectivorous bats".Acta Chiropterologica.9(1): 261–267.doi:10.3161/1733-5329(2007)9[261:COGPBF]2.0.CO;2.S2CID86038700.
  44. ^abTamisiea, Jack (November 5, 2024)."You May Not Be Able to Outrun a Vampire Bat".The New York Times.
  45. ^Schneider, Maria Cristina; Romijn, Phyllis Catharina; Uieda, Wilson; Tamayo, Hugo; Silva, Daniela Fernandes da; Belotto, Albino; Silva, Jarbas Barbosa da; Leanes, Luis Fernando (March 2009)."Rabies transmitted by vampire bats to humans: an emerging zoonotic disease in Latin America?".Revista Panamericana de Salud Pública.25(3): 260–269.doi:10.1590/S1020-49892009000300010.PMID19454154.
  46. ^"The Art and Science of Bats".Smithsonian Institution.
  47. ^Davis, April; Gordy, Paul; Rudd, Robert; Jarvis, Jodie A.; Bowen, Richard A. (2012)."Naturally Acquired Rabies Virus Infections in Wild-Caught Bats".Vector-Borne and Zoonotic Diseases.12(1): 55–60.doi:10.1089/vbz.2011.0674.ISSN1530-3667.PMC3249890.PMID21923271.
  48. ^"Rabies in bats: how to spot it and report it - Signs that a bat may have rabies".UK Government - Environment.19 January 2023.
  49. ^Hacke, Werner; Albers, Greg; Al-Rawi, Yasir; Bogousslavsky, Julien; Davalos, Antonio; Eliasziw, Michael; Fischer, Michael; Furlan, Anthony; Kaste, Markku; Lees, Kennedy R.; Soehngen, Mariola; Warach, Steven (2005)."The Desmoteplase in Acute Ischemic Stroke Trial (DIAS)".Stroke.36(1): 66–73.doi:10.1161/01.str.0000149938.08731.2c.ISSN0039-2499.PMID15569863.A search for "desmoteplase site:ahajournals.org" will find other studies in American Heart Association journals.

Further reading

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Wikimedia Commons has media related toDesmodontinae.
  • Greenhall, A., G. Joermann, U. Schmidt, M. Seidel. 1983. Mammalian Species: Desmodus rotundus. American Society of Mammalogists, 202: 1–6.
  • Campbell A, Naik RR, Sowards L, Stone M (2002)."Biological infrared imaging and sensing"(PDF).Micron.33(2): 211–225.doi:10.1016/S0968-4328(01)00010-5.PMID11567889.Archived fromthe original(PDF)on 2003-06-15.
  • Pawan, J.L. (1936b). "Rabies in the Vampire Bat of Trinidad with Special Reference to the Clinical Course and the Latency of Infection."Annals of Tropical Medicine and Parasitology.Vol. 30, No. 4. December, 1936.
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