If you have trouble with accessing the paper, data, or R code, please email me.
See also Google Scholar.

Submitted manuscripts

Crisp R, Brent L, and Carter G. Social dominance and cooperation in female vampire bats. (preprint)

Hobson L and Carter G. Network approaches to understanding social organization and complexity [book chapter in Handbook of Comparative Psychology, Edited by Amanda Ridley, Patrizia d’Ettorre, and Todd Freeberg] (draft)

Farine D and Carter G. Permutation tests for hypothesis testing with animal social network data: problems and potential solutions. (preprint).

Brown BKG and Carter GG. Do bats use scent cues to find roosts? [review paper] (preprint)


Wilkinson GS, Adams DM, Arnold BD, Ball HC, Breeze CE, Carter GG, Cooper LN et al. In press. Genome methylation predicts age and longevity of bats. Nature Communications (preprint).

Carter GG. In press. Species spotlight: Co-option and the evolution of food sharing in vampire bats. Ethology (preprint)

Razik I, Brown BKG, Page RA, Carter GG. 2021. Nonkin adoption in the common vampire bat. Royal Society Open Science. 8: 201927. link.

Cantor M, Maldonado-Chaparro AA, Beck KB, Brandl HB, Carter GG, He P, Hillemann F, Klarevas-Irby JA, Ogino M, Papageorgiou D, Prox L, Farine DR. 2021. The importance of individual-to-society feedbacks in animal ecology and evolution. Journal of Animal Ecology. 90: 27-44. (link)

Carter G, Brown B, Razik, I, Ripperger S. 2021. Penguins, falcons, and mountain lions: the extraordinary host diversity of vampire bats. In 50 Years of Bat Research: Foundations and New Frontiers. Springer. pp 151-170. (link) (PDF). [book chapter]


Ripperger S, Stockmaier B, Carter G. 2020. Tracking sickness effects on social encounters via continuous proximity-sensing in wild vampire bats. Behavioral Ecology. 31:1296–1302. link

Brown B, Leffer L, Valverde Y, Toshkova N, Nystrom J, Page RA, Carter G. 2020. Do bats use guano and urine stains to find new roosts? Tests with three group-living bats. Royal Society Open Science. 7:201055.

Carter G, Farine D, Crisp R, Vrtilek J, Ripperger S, Page R. 2020. Development of new food-sharing relationships in vampire bats. Current Biology. 30(7): 1275-1279.e3. (PDF).

Stockmaier S, Bolnick DI, Page RA, Josic D, Carter G. 2020. Immune-challenged vampire bats produce fewer contact calls. Biology Letters. (PDF)
Press: New York Times (download_pdf)

Kohles J, Carter G, Page R, Dechmann D. 2020. The social foraging bat Molossus molossus discriminates between individual group members based on their search-phase echolocation calls. Behavioral Ecology. araa056. (PDF)

Stockmaier S, Bolnick D, Page R, Carter G. 2020. Sickness effects on social interactions depend on the type of behaviour and relationship. Journal of Animal Ecology. 89: 1387-1394. (PDF)

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. 2020. Thinking small: next-generation sensor networks close the size gap in vertebrate biologging. PLoS Biology. 18(4): e3000655.

Ripperger SP, Duda N, Kölpin A, Carter G. 2020. Simultaneous monitoring of the same animals with PIT-tags and sensor nodes causes no system interference. Animal Behavior and Cognition. 7(4), 531-536. (link)

Geipel* I, Kernan* CE, Litterer AS, Carter GG, Page RA, ter Hofstede HM. 2020. Predation risks of signalling and searching: bats prefer katydids in motion. Biology Letters. 16:20190837.

Flores V*, Carter G*, Halczok T, Kerth G, Page R. 2020. Social structure and relatedness in the fringe-lipped bat (Trachops cirrhosus). Royal Society Open Science. 7:192256.

Narizano H, Carter GG. 2020. Do vampire bats groom others based on need? Behavioral Ecology. 31:107-113 (PDF)

Carter G, Chen T, Razik I. 2020. The Theory of Reciprocal Altruism. In: The SAGE Handbook of Evolutionary Psychology (editor: T Shackelford). Sage. [PDF]

Hermanson J, Carter G. 2020. 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* (*equal contributions), 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. 29: 4139-4144.e4. (PDF)

Carter G, Schino G, Farine D. 2019. Challenges with assessing the roles of nepotism and reciprocity in cooperation networks. Animal Behavior. 150:255-271 (PDF)

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. (PDF)

Gould GG, Carter GG, Augustine JK. 2019. Divergent signals from homologous unfeathered ornaments in two congeneric grouse. Ecology and Evolution. 9: 11833-11847.

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

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.


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. 5:172483.

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.

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

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;

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


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

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

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


Carter G, Wilkinson G. 2015. Social benefits of non-kin food sharing by female vampire bats. Proceedings of the Royal Society B. 282: 20152524.

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.

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.

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

2014 and earlier

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

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.

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.

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.

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. (“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.

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

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-peer-reviewed publications

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

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

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