In review

Ripperger SP, Page RA, Mayer F, Carter GG. Evidence for unfamiliar kin recognition in vampire bats. (preprint) Submitted to Biology Letters.

Kohles J, Carter GG, Page R, Dechmann D. The social foraging bat Molossus molossus discriminates between individual group members based on their search-phase echolocation calls. (email us for preprint) In review at Behavioral Ecology.

Geipel* I, Kernan* CE, Litterer AS, Carter GG, Page RA, ter Hofstede HM. Predation risks of signalling and searching: bats prefer katydids in motion. (email us for preprint) In revision for Biology Letters.

Ripperger SP, Carter GG, Page RA, Duda N, Koelpin A, Weigel R, Hartmann M, Nowak T, Thielecke J, Schadhauser M, Robert J, Herbst S, Meyer-Wegener K, Wägemann P, Schröder-Preikschat W, Cassens B, Kapitza R, Dressler F, Mayer F. Thinking small: next-generation sensor networks close the size gap in vertebrate biologging. (preprint). Provisional acceptance with minor revisions at PLoS Biology.


Carter G, Farine D, Crisp R, Vrtilek J, Ripperger S, Page R. Development of new food-sharing relationships in vampire bats. (preprint). In press. Current Biology.

Carter G, Brown B, Razik, I, Ripperger S. Penguins, falcons, and mountain lions: the extraordinary host diversity of vampire bats. In 50 Years of Bat Research. Foundations and New Frontiers: Fascinating Life Sciences. Springer. (preprint). In press.

Stockmaier S, Bolnick D, Page R, Carter G. Effects of sickness on social networks depend on the type of behaviour, measure, and relationship. In press (email us for preprint). Journal of Animal Ecology.

Carter G, Chen T, Razik I. In press. The Theory of Reciprocal Altruism. In: The SAGE Handbook of Evolutionary Psychology (editor: T Shackelford). Sage. (preprint) (book chapter)

Hermanson J & Carter G. In press. Vampire bats. In: Phyllostomid Bats, a Unique Mammalian Radiation (editors: TH Fleming, L Davalos, M Mello). University of Chicago Press. (preprint)(book chapter)


Ripperger SP*, Carter GG*, Duda N, Koelpin A, Cassens B, Kapitza R, Josic D, Berrío-Martínez J, Page RA, Mayer F. Vampire bats that cooperate in the lab maintain their social networks in the wild. 2019. Current Biology. (*equal contributions). (link) Press: PBSCNNNPR , BBCNature MagazineScience MagazineScience NewsPopular ScienceThe Ohio State UniversityCosmos MagazineWissenschaftEl MundoZME ScienceSciShow, EurekaAlert Press release (Attention received in top 1% of all research papers of the same age, as scored by Altmetric, in Top 5 “Coolest Science Stories” at OSU)

Narizano H, Carter GG. 2019. Do vampire bats groom others based on need? Behavioral Ecology. (PDF)

Gould GG, Carter GG, Augustine JK. 2019. Divergent signals from homologous unfeathered ornaments in two congeneric grouse. Ecology and Evolution.

Berrío-Martínez J, Kaiser S, Nowak M, Page R, Carter G. 2019. The role of past experience in the development of feeding behavior in common vampire bats. PeerJ. 7:e7448

Carter G, Schino G, and Farine D. 2019. Challenges with assessing the roles of nepotism and reciprocity in cooperation networks. Animal Behavior. 149:1-17 (PDF)

Vleut I, Carter G, Medellin R. 2019. Movement ecology of the carnivorous woolly false vampire bat (Chrotopterus auritus) in southern Mexico. PLoS ONE. 14(7):e0220504.

Wilkinson G, G Carter, KM Bohn, B Caspers, G Chaverri, D Farine, L Günther, G Kerth, M Knörnschild, F Mayer, M Nagy, J Ortega & K Patriquin. 2019. Kinship, association and social complexity in bats. Behavioral Ecology and Sociobiology. 73: 7. preprint PDF


Stockmaier S, Bolnick DI, Page RA, Carter GG. 2018. An immune challenge reduces social grooming in vampire bats. Animal Behaviour 140: 141-149.(PDF

Vrtilek J, Carter G, Patriquin K, Page R and J Ratcliffe. 2018. A method for rapid testing of social learning in vampire bats. Royal Society Open Science. DOI: 10.1098/rsos.172483

Carter GG, Forss S, Page RA, Ratcliffe JM. 2018. Younger vampire bats (Desmodus rotundus) are more likely than adults to explore novel objects. PLoS ONE 13(5): e0196889.


Carter G, Farine D, Wilkinson G. 2017. Social bet-hedging in vampire bats. Biology Letters. 13:20170112. doi: 10.1098/rsbl.2017.0112 (link)(text)(PDF)

Carter G, Wilkinson G, Page R. 2017. Food-sharing vampire bats are more nepotistic under conditions of perceived risk. Behavioral Ecology. 28: 565-569. (PDF) (data)

Carter G. 2017. Gerald S. Wilkinson. In: Vonk J & Shackelford T (eds). Encyclopedia of Animal Cognition and Behavior. (link) (PDF)

Delpietro HA, Russo R, Carter G, Lord R, Delpietro G. 2017. Reproductive seasonality, sex ratio, and philopatry in Argentina’s common vampire bats. Royal Society Open Science. 4:160959; DOI: 10.1098/rsos.160959.  (open access)


Carter G, Wilkinson G. 2016. Common vampire bat contact calls attract past food-sharing partners. Animal Behavior. 116:45-51. (PDF) (data)

Wilkinson G, Carter G, Bohn K, Adams D. 2016. Non-kin cooperation in bats. Philosophical Transactions B. doi: 10.1098/rstb.2015.0095 (PDF)

Carter G. 2016. Blood sharing in vampire bats. In: Shackelford TK, Weekes-Shackelford V (eds). Encyclopedia of Evolutionary Psychological Science. (PDF)

Carter G, Bohn K. 2016. “Altruism” in nonkin. In: Shackelford TK, Weekes-Shackelford V (eds). Encyclopedia of Evolutionary Psychological Science. (link)

Bohn K, Carter G. 2016. Reciprocal Altruism and Cooperation for Mutual Benefit. In: Shackelford TK, Weekes-Shackelford V (eds). Encyclopedia of Evolutionary Psychological Science. (link)

Carter G. 2016. Nonhuman reciprocal altruism. In: Shackelford TK, Weekes-Shackelford V (eds). Encyclopedia of Evolutionary Psychological Science. (link)

2015 and earlier (before PhD)

Carter G, Wilkinson G. 2015. Social benefits of non-kin food sharing by female vampire bats. Proceedings of the Royal Society B. 282: 20152524. (PDF) (data)(supplement) press: SapiensNational Geographic,, Science News, Broadly

Carter G, Wilkinson G. 2015. Intranasal oxytocin increases social grooming and food sharing in the common vampire bat. Hormones and Behavior. 75: 150-153. (PDF) press: Newsweek

Carter G, Leffer L. 2015. Social grooming in bats: are vampire bats exceptional? PLOS One. 10(10):e0138430. doi: 10.1371/journal.pone.0138430. (open access) (data) press: National Geographic,  NewStatesmanThe Australian,  iScience,  Nature World News

Carter G, Stewart A. 2015. The floral bat lure dimethyl disulphide does not attract the palaeotropical Dawn bat. Journal of Pollination Ecology. 17: 129-131. (open access)

Carter G, Schoeppler D, Manthey M, Knoernschild M, Denzinger A. 2015. Distress calls of a fast-flying aerial-hawking bat (Molossus molossus) provoke inspection flights but not cooperative mobbing. PLOS One. 10(9): e0136146. doi:10.1371/journal.pone.0136146  (open access) (data)

Carter, G. 2014. The reciprocity controversy. Animal Behavior and Cognition. 1(3), 368-386. doi:10.12966/abc.08.11.2014 (open access)

Kershenbaum A, D Blumstein, M Roch, C Akcay, G Backus, M Bee, K Bohn, Y Cao, G Carter et al. (42 authors) 2014. Acoustic sequences in non-human animals: A tutorial review and prospectus. Biological Reviews. doi:10.1111/brv.12160 (PDF)

Carter G, Wilkinson G. 2013. Does food sharing in vampire bats demonstrate reciprocity? Communicative & Integrative Biology. 6:e25783. (open access)

Carter G, Wilkinson G. 2013. Cooperation and conflict in the social lives of bats. In: Adams, R, Pedersen, S (eds). Bat Evolution, Ecology, and Conservation. Springer Science Press. pg 225-242. (PDF) (e-book)

Carter G, Wilkinson G. 2013. Food sharing in vampire bats: reciprocal help predicts donations more than relatedness or harassment. Proceedings of the Royal Society B. 280: 1753. doi: 10.1098/rspb.2012.2573 (open access, text supplement, videoposter, data) press: Current Biology, BBCEarth Sky, Science News

Carter G, R Logsdon, B Arnold, A Menchaca, R Medellin. 2012. Adult vampire bats produce contact calls when isolated: acoustic variation between species, colonies, and individuals. PLOS One. 7(6): e38791. doi:10.1371/journal.pone.0038791. (open access, “top 25% most cited PLOS ONE articles”)

Carter G, J Ratcliffe, and B Galef. 2010. Flower bats (Glossophaga soricina) and fruit bats (Carollia perspicillata) rely on spatial cues over shapes and scents when relocating food. PLOS One. 5(5): e10808. (open access)

Carter G, B Fenton, and P Faure. 2009. White-winged vampire bats (Diaemus youngi) exchange contact calls. Canadian Journal of Zoology. 87:604-608. (PDF)

Carter G, M Skowronski, P Faure, B Fenton. 2008. Antiphonal calling allows individual discrimination in white-winged vampire bats. Animal Behaviour. 76:1343-1355. (PDF) (Video supplement) press: Discovery News

Riskin D, S Parsons, W Schutt, G Carter, J Hermanson. 2006. Terrestrial locomotion of the New Zealand short-tailed bat (Mystacina tuberculata) and the common vampire bat (Desmodus rotundus). Journal of Experimental Biology. 209:1725-1736. (PDF) press: Slate, JEB

Carter G, D Riskin. 2006. Mystacina tuberculataMammalian Species. 790:1-8. (PDF)

Carter G, C Coen, L Stenzler, I Lovette. 2006. Avian host DNA isolated from the feces of white-winged vampire bats (Diaemus youngi)Acta Chiropterologica. 8(1):255-259. (PDF) press: Living Bird

Non-refereed publications, data, code and media 

Carter, G. 2018. Do vampire bats have friends? Bat News. PDF

Carter, G. 2018. supplementary R code for Wilkinson et al. “Kinship, association and social complexity in bats” in Behavioral Ecology and Sociobiology. Figshare

Carter, G. 2016. Footage of feeding by the hairy-legged vampire bat Diphylla ecaudata refutes the hypothesis that this vampire bat sucks. Figshare.

Carter, G. 2016. Footage of regurgitated food sharing among adult common vampire bats. Figshare.

Carter, G. 2015. Cooperation and social bonds in common vampire bats. PhD Thesis. University of Maryland.

Carter G. 2011. The future of bat research and conservation. Sonorensis Magazine: Celebrating Bats. Arizona-Sonora Desert Museum. 31:1. (PDF)

Carter G. 2011. Food sharing in vampire bats. Bat Conservation Journal. Organization for Bat Conservation. Fall 2011 Issue.

Carter G. 2005. Bat diversity and abundance in Cusuco National Park core zone, Honduras. Internal report for Operation Wallacea, UK. (PDF)