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Professor Darren Croft

Professor Darren Croft

Professor of Behavioural Ecology


 Washington Singer 122


Washington Singer Laboratories, University of Exeter, Perry Road, Prince of Wales Road, Exeter, EX4 4QG, UK


I am a behavioural ecologist and my research interests lie in two main questions (i) What are the mechanisms and functions that underpin the structure of animal societies? (ii) What are the ecological and evolutionary implications of this structure? To address these questions my research group combines experimental and observational work on wild animal populations with controlled laboratory experiments. The research topics covered by my group are broad and include; the evolution of cooperation, life history evolution, social recognition and sexual conflict. As such we work on a wide range of study systems ranging from small freshwater fish to killer whales. 

I have recently been appointed as Executive Director of the Center for Whale Research in the USA, which is responsible for collecting one of the longest and most complete individual-based studies of a killer whale population anywhere in the world. I have been collaborating with the Centre for Whale Research for over a decade on research focused on the sociobiology and conservation of killer whales. Killer whales are one of just 6 species of mammals that go through menopause and working with the long-term data collected by the Center for Whale Research, we have examined how and why this unusual life history trait evolved. One of our study populations (the southern resident killer whales) is also critically endangered and our research in partnership with the Center for Whale Research is examining how social factors contribute to survival and reproduction, essential information needed to inform future management and conservation strategies. 

In the news: Listen to Darren talking about the role of grandmothers and his research on the evolution of menopause in killer whales on this BBC Radio 4 documentary or learn about his work in this TedEd video that has been watched over 1 million times. 

Research Keywords:

  • Comparative psychology
  • Group living
  • Cooperation 
  • Social evolution
  • Ageing 
  • Health and welfare


2017: Fellow of the Higher Education Academy

2000-2004: PhD in Behavioural Ecology, University of Leeds.

1996-1999: Bachelor of Science with Honours, Biology (Class I), University of Leeds.


2016-Current: Professor of Animal Behaviour, University of Exeter

2014-2016: Associate Professor of Animal Behaviour, University of Exeter

2009-2014: Senior Lecturer in Animal Behaviour, University of Exeter

2008-2009: Lecturer in Animal Behaviour, University of Exeter

2006-2008: Lecturer in Animal Behaviour, University of Wales Bangor

2004-2006: Post Doctoral Research Fellow, University of Leeds


Research group links

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Research projects

Broadly my research can be divided into two distinct areas that cross-fertilise one another:

i) The ecology and evolution of group living in wild animal populations: This work combines experimental and observational work in natural populations together with controlled laboratory manipulations and experiments to provide a very powerful framework for testing behavioural and evolutionary hypotheses.

ii) The implications of social behaviour for the management, welfare and productivity of captive animals. Building on my work on wild animal populations, my research group is applying methods and concepts that have been developed in behavioural ecology to real world questions relating to animal health, welfare and productivity.

I welcome informal applications from potential PhD students and post-doctoral fellows who are able to attract their own funding (e.g. international PhD scholarships, EU fellowships and visiting researchers). If you are interested in joining our research group then please email me.

Research grants

  • 2019 NERC
    The Evolution of Sex Differences in Mammalian Social Life Histories
  • 2019 NERC
    Leaders of war: the evolution of collective decision-making in the face of intergroup conflict
  • 2018 NERC
    The ecology and evolution of intergroup conflict in animal societies: theory and tests
  • 2016 European Research Council
    Evolutionary genetics of guppy sex chromosomes. Led by PI: Prof Deborah Charlesworth, University of Edinburgh
  • 2015 Leverhulme Trust
    The Evolution of Eye Salience as a Signal for Communication
  • 2015 NERC
    Infection In Dynamic Social Networks of a Wild Mammal
  • 2014 Pets at Home
    Stress and Mortality in Aquarium Fish: Interventions for Improving Health and Well-being
  • 2013 NERC
    The Evolution of Prolonged Post-Reproductive Lifespan in a Non-Human Mammal
  • 2013 FNU (The Danish Council for Independent Research / Natural S
    Social niche construction and evolutionary implications for animal behaviour
  • 2012 BBSRC
    Assessment of Dairy Cow Welfare through Predictive Modelling of Individual and Social Behaviour
  • 2011 Leverhulme Trust
    The Evolution of Cooperation in Structured Animal Societies.
  • 2011 DairyCo
    Social Components of Welfare and Productively in Dairy Cattle
  • 2011 DEFRA
    The influence of Social Networks on Welfare and Productivity in Dairy Cattle
  • 2010 FNU (The Danish Council for Independent Research)
    Cognitive and Physical Aspects of Animal Communication Networks
  • 2007 Nuffield Science Award
    Behavioural Syndromes: Environmental Influences in a Naturally Clonal Species of Fish
  • 2006 NERC
    Mechanisms and Functions of Sexual Segregation
  • 2004 Fisheries Society of the British Isles
    Sexual Segregation in the Trinidadian Guppy

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Krause J, James R, Franks D, Croft D (2014). Animal Social Networks., Oxford University Press. Abstract.
Croft DP, James R, Krause J (2008). Exploring Animal Social Networks. Princetown, NJ, Princetown University Press.

Journal articles

Langford J, Fawcett T, Price E, Bradley D, Wilson A, Croft D (In Press). Bio-logging reveals heritable patterns of natural behaviours in sheep. Proceedings of 2023 IEEE International Workshop on Measurements and Applications in Veterinary and Animal Sciences, IEEE Catalog Number CFP23CX4-USB, 182-187.
Chopra K, Hodges HR, Barker ZE, Vázquez Diosdado JA, Amory JR, Cameron TC, Croft DP, Bell NJ, Thurman A, Bartlett D, et al (2024). Bunching behavior in housed dairy cows at higher ambient temperatures. J Dairy Sci, 107(4), 2406-2425. Abstract.  Author URL.
Ellis S, Franks DW, Nielsen MLK, Weiss MN, Croft DP (2024). The evolution of menopause in toothed whales. Nature, 627(8004), 579-585. Abstract.  Author URL.
Dimitriadou S, Winberg S, Thörnqvist PO, Croft DP, Darden SK (2023). Brain monoaminergic activity during predator inspection in female Trinidadian guppies (Poecilia reticulata). Behavioural Brain Research, 436 Abstract.
Weiss MN, Ellis S, Franks DW, Nielsen MLK, Cant MA, Johnstone RA, Ellifrit DK, Balcomb KC, Croft DP (2023). Costly lifetime maternal investment in killer whales. Current Biology, 33(4), 744-748.e3.
Weiss MN, Croft DP (2023). Killer whales. Current Biology, 33(12), r668-r670.
Grimes C, Brent LJN, Ellis S, Weiss MN, Franks DW, Ellifrit DK, Croft DP (2023). Postreproductive female killer whales reduce socially inflicted injuries in their male offspring. Current Biology, 33(15), 3250-3256.e4.
Lewis KE, Price E, Croft DP, Green LE, Ozella L, Cattuto C, Langford J (2023). Potential role of biologgers to automate detection of lame ewes and lambs. Applied Animal Behaviour Science, 259 Abstract.
Lybkær Kronborg Nielsen M, Ellis S, Weiss MN, Towers JR, Doniol-Valcroze T, Franks DW, Cant MA, Ellis GM, Ford JKB, Malleson M, et al (2023). Temporal dynamics of mother-offspring relationships in Bigg’s killer whales: opportunities for kin-directed help by post-reproductive females. Proceedings of the Royal Society B: Biological Sciences
Ozella L, Price E, Langford J, Lewis KE, Cattuto C, Croft DP (2022). Association networks and social temporal dynamics in ewes and lambs. Applied Animal Behaviour Science, 246 Abstract.
Price E, Langford J, Fawcett TW, Wilson AJ, Croft DP (2022). Classifying the posture and activity of ewes and lambs using accelerometers and machine learning on a commercial flock. Applied Animal Behaviour Science, 251 Abstract.
Jacoby DMP, Brown C, Croft DP, Mann J, Mourier J (2022). Editorial: Sociality in the Marine Environment. Frontiers in Marine Science, 9
Sankey DWE, Hunt KL, Croft DP, Franks DW, Green PA, Thompson FJ, Johnstone RA, Cant MA (2022). Leaders of war: Modelling the evolution of conflict among heterogeneous groups. Philosophical Transactions of the Royal Society B: Biological Sciences, 377(1851). Abstract.
Qiu S, Yong L, Wilson A, Croft DP, Graham C, Charlesworth D (2022). Partial sex linkage and linkage disequilibrium on the guppy sex chromosome. Molecular Ecology, 31(21), 5524-5537. Abstract.
Ellis S, Johnstone RA, Cant MA, Franks DW, Weiss MN, Alberts SC, Balcomb KC, Benton CH, Brent LJN, Crockford C, et al (2022). Patterns and consequences of age-linked change in local relatedness in animal societies. Nature Ecology and Evolution, 6(11), 1766-1776. Abstract.
Yong L, Croft DP, Troscianko J, Ramnarine IW, Wilson AJ (2022). Sensory-based quantification of male colour patterns in Trinidadian guppies reveals no support for parallel phenotypic evolution in multivariate trait space. Mol Ecol, 31(5), 1337-1357. Abstract.  Author URL.
Lewis KE, Price E, Croft DP, Langford J, Ozella L, Cattuto C, Green LE (2022). Social behaviour and transmission of lameness in a flock of ewes and lambs. Frontiers in Veterinary Science, 9 Abstract.
Dimitriadou S, Santos EM, Croft DP, van Aerle R, Ramnarine IW, Filby AL, Darden SK (2022). Social partner cooperativeness influences brain oxytocin transcription in Trinidadian guppies (Poecilia reticulata). Behavioural Brain Research, 423, 113643-113643.
Dimitriadou S, Santos EM, Croft DP, van Aerle R, Ramnarine IW, Filby AL, Darden SK (2022). Social partner cooperativeness influences brain oxytocin transcription in Trinidadian guppies (Poecilia reticulata). Behavioural Brain Research, 423 Abstract.
Grimes C, Brent LJN, Weiss MN, Franks DW, Balcomb KC, Ellifrit DK, Ellis S, Croft DP (2022). The effect of age, sex, and resource abundance on patterns of rake markings in resident killer whales (Orcinus orca). Marine Mammal Science, 38(3), 941-958. Abstract.
Nielsen MLK, Ellis S, Towers JR, Doniol-Valcroze T, Franks DW, Cant MA, Weiss MN, Johnstone RA, Balcomb KC, Ellifrit DK, et al (2021). A long postreproductive life span is a shared trait among genetically distinct killer whale populations. Ecology and Evolution, 11(13), 9123-9136. Abstract.
Weiss MN, Franks DW, Giles DA, Youngstrom S, Wasser SK, Balcomb KC, Ellifrit DK, Domenici P, Cant MA, Ellis S, et al (2021). Age and sex influence social interactions, but not associations, within a killer whale pod. Proceedings of the Royal Society B: Biological Sciences, 288(1953). Abstract.
Brask JB, Ellis S, Croft DP (2021). Animal social networks: an introduction for complex systems scientists. Journal of Complex Networks, 9(2), 1-19. Abstract.
Green PA, Preston EFR, Nicholl MH, Croft DP, Thompson FJ, Cant MA (2021). Collective defence and behavioural homogeneity during simulated territorial intrusions in banded mongooses (<i>Mungos mungo</i>). ETHOLOGY, 127(10), 886-896.  Author URL.
Weiss MN, Franks DW, Brent LJN, Ellis S, Silk MJ, Croft DP (2021). Common datastream permutations of animal social network data are not appropriate for hypothesis testing using regression models. Methods in Ecology and Evolution, 12(2), 255-265. Abstract.
Weiss MN, Ellis S, Croft DP (2021). Diversity and Consequences of Social Network Structure in Toothed Whales. Frontiers in Marine Science, 8 Abstract.
Allen CRB, Brent LJN, Motsentwa T, Croft DP (2021). Field evidence supporting monitoring of chemical information on pathways by male African elephants. Animal Behaviour, 176, 193-206. Abstract.
Allen CRB, Croft DP, Testard C, Brent LJN (2021). Function of trunk-mediated “greeting” behaviours between male african elephants: Insights from choice of partners. Animals, 11(9). Abstract.
Macario A, Darden SK, Verbruggen F, Croft DP (2021). Intraspecific variation in inhibitory motor control in guppies, Poecilia reticulata. Journal of Fish Biology, 98(1), 317-328. Abstract.
Croft DP, Weiss MN, Nielsen MLK, Grimes C, Cant MA, Ellis S, Franks DW, Johnstone RA (2021). Kinship dynamics: patterns and consequences of changes in local relatedness. Proceedings of the Royal Society B: Biological Sciences, 288(1957), 20211129-20211129. Abstract.
Ellis S, Franks DW, Weiss MN, Cant MA, Domenici P, Balcomb KC, Ellifrit DK, Croft DP (2021). Mixture models as a method for comparative sociality: social networks and demographic change in resident killer whales. Behavioral Ecology and Sociobiology, 75(4). Abstract.
Preston EFR, Thompson FJ, Ellis S, Kyambulima S, Croft DP, Cant MA (2021). Network-level consequences of outgroup threats in banded mongooses: Grooming and aggression between the sexes. Journal of Animal Ecology, 90(1), 153-167. Abstract.
Allen CRB, Croft DP, Brent LJN (2021). Reduced older male presence linked to increased rates of aggression to non-conspecific targets in male elephants. Proceedings of the Royal Society B: Biological Sciences, 288(1965). Abstract.
Preston EFR, Thompson FJ, Kyabulima S, Croft DP, Cant MA (2021). The dynamics of social cohesion in response to simulated intergroup conflict in banded mongooses. ECOLOGY AND EVOLUTION, 11(24), 18662-18675.  Author URL.
Heathcote RJP, Troscianko J, Darden S, Naisbett-Jones LC, Laker P, Brown AM, Ramnarine IW, Walker J, Croft D (2020). A matador-like predator diversion strategy driven by conspicuous colouration in guppies. Current Biology, 30 (14), 2844-2851.
Franks DW, Weiss MN, Silk MJ, Perryman RJY, Croft DP (2020). Calculating effect sizes in animal social network analysis. Methods in Ecology and Evolution, 12(1), 33-41. Abstract.
Rose PE, Croft DP (2020). Evaluating the social networks of four flocks of captive flamingos over a five-year period: Temporal, environmental, group and health influences on assortment. Behavioural Processes, 175 Abstract.
Allen CRB, Brent LJN, Motsentwa T, Weiss MN, Croft DP (2020). Importance of old bulls: leaders and followers in collective movements of all-male groups in African savannah elephants (Loxodonta africana). Scientific Reports, 10(1). Abstract.
Weiss MN, Franks DW, Balcomb KC, Ellifrit DK, Silk MJ, Cant MA, Croft DP (2020). Modelling cetacean morbillivirus outbreaks in an endangered killer whale population. Biological Conservation, 242 Abstract.
Chopra K, Hodges HR, Barker ZE, Vazquez Diosdado JA, Amory JR, Cameron TC, Croft DP, Bell NJ, Codling EA (2020). Proximity Interactions in a Permanently Housed Dairy Herd: Network Structure, Consistency, and Individual Differences. FRONTIERS IN VETERINARY SCIENCE, 7  Author URL.
Ozella L, Langford J, Gauvin L, Price E, Cattuto C, Croft DP (2020). The effect of age, environment and management on social contact patterns in sheep. APPLIED ANIMAL BEHAVIOUR SCIENCE, 225  Author URL.
Darden SK, James R, Cave JM, Brask JB, Croft DP (2020). Trinidadian guppies use a social heuristic that can support cooperation among non-kin. Proceedings. Biological sciences, 287(1934). Abstract.
Dimitriadou S, Croft DP, Darden SK (2019). Divergence in social traits in Trinidadian guppies selectively bred for high and low leadership in a cooperative context. Scientific Reports, 9(1). Abstract.
Brask JB, Croft DP, Edenbrow M, James R, Bleakley BH, Ramnarine IW, Heathcote RJP, Tyler CR, Hamilton PB, Dabelsteen T, et al (2019). Evolution of non-kin cooperation: social assortment by cooperative phenotype in guppies. R Soc Open Sci, 6(1). Abstract.  Author URL.
Cant MA, Croft DP (2019). Grandmothering in space and time. Current Biology
Silk MJ, Hodgson DJ, Rozins C, Croft DP, Delahay RJ, Boots M, McDonald RA (2019). Integrating social behaviour, demography and disease dynamics in network models: Applications to disease management in eclining wildlife populations. Philosophical Transactions of the Royal Society B: Biological Sciences, 374(1781). Abstract.
Macario A, Croft DP, Darden SK (2019). Male phenotypic diversity experienced during ontogeny mediates female mate choice in guppies. Behavioral Ecology, 30(2), 465-473. Abstract.
Weiss MN, Franks DW, Croft DP, Whitehead H (2019). Measuring the complexity of social associations using mixture models. BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY, 73(1).  Author URL.
Nattrass S, Croft DP, Ellis S, Cant MA, Weiss MN, Wright BM, Stredulinsky E, Doniol-Valcroze T, Ford JKB, Balcomb KC, et al (2019). Postreproductive killer whale grandmothers improve the survival of their grandoffspring. Proceedings of the National Academy of Sciences of the United States of America, 116(52), 26669-26673. Abstract.
Ellis S, Franks DW, Nattrass S, Currie TE, Cant MA, Giles D, Balcomb KC, Croft DP (2018). Analyses of ovarian activity reveal repeated evolution of post-reproductive lifespans in toothed whales. SCIENTIFIC REPORTS, 8  Author URL.
Silk MJ, Weber NL, Steward LC, Hodgson DJ, Boots M, Croft DP, Delahay RJ, McDonald RA (2018). Contact networks structured by sex underpin sex-specific epidemiology of infection. Ecology Letters, 21(2), 309-318. Abstract.
Heathcote RJP, Darden SK, Troscianko J, Lawson MRM, Brown AM, Laker PR, Naisbett-Jones LC, MacGregor HEA, Ramnarine I, Croft DP, et al (2018). Dynamic eye colour as an honest signal of aggression. Current Biology, 28(11), R652-R653.
Rose PE, Brereton JE, Croft DP (2018). Measuring welfare in captive flamingos: Activity patterns and exhibit usage in zoo-housed birds. Applied Animal Behaviour Science, 205, 115-125. Abstract.
Rose PE, Lloyd I, Brereton JE, Croft DP (2018). Patterns of nocturnal activity in captive greater flamingos. Zoo Biology, 37(5), 290-299. Abstract.
Ellis S, Franks DW, Nattrass S, Cant MA, Bradley DL, Giles D, Balcomb KC, Croft DP (2018). Postreproductive lifespans are rare in mammals. Ecology and Evolution, 8(5), 2482-2494. Abstract.
Silk MJ, Drewe JA, Delahay RJ, Weber N, Steward LC, Wilson-Aggarwal J, Boots M, Hodgson DJ, Croft DP, McDonald RA, et al (2018). Quantifying direct and indirect contacts for the potential transmission of infection between species using a multilayer contact network. Behaviour, 155(7-9), 731-757. Abstract.
Rose PE, Croft DP (2018). Quantifying the social structure of a large captive flock of greater flamingos (Phoenicopterus roseus): Potential implications for management in captivity. Behavioural Processes, 150, 66-74. Abstract.
Rozins C, Silk MJ, Croft DP, Delahay RJ, Hodgson DJ, McDonald RA, Weber N, Boots M (2018). Social structure contains epidemics and regulates individual roles in disease transmission in a group-living mammal. Ecology and Evolution, 8(23), 12044-12055. Abstract.
Diosdado JAV, Barker ZE, Hodges HR, Amory JR, Croft DP, Bell NJ, Codling EA (2018). Space-use patterns highlight behavioural differences linked to lameness, parity, and days in milk in barn-housed dairy cows. PLOS ONE, 13(12).  Author URL.
Barker ZE, Diosdado JAV, Codling EA, Bell NJ, Hodges HR, Croft DP, Amory JR (2018). Use of novel sensors combining local positioning and acceleration to measure feeding behavior differences associated with lameness in dairy cattle. JOURNAL OF DAIRY SCIENCE, 101(7), 6310-6321.  Author URL.
Macario A, Croft DP, Endler JA, Darden SK (2017). Early social experience shapes female mate choice in guppies. Behavioral Ecology, 28(3), 833-843. Abstract.
Lacy RC, Williams R, Ashe E, Balcomb KCIII, Brent LJN, Clark CW, Croft DP, Giles DA, MacDuffee M, Paquet PC, et al (2017). Evaluating anthropogenic threats to endangered killer whales to inform effective recovery plans. SCIENTIFIC REPORTS, 7  Author URL.
Heathcote RJP, Darden SK, Franks DW, Ramnarine IW, Croft DP (2017). Fear of predation drives stable and differentiated social relationships in guppies. Scientific Reports, 7 Abstract.
Ellis S, Franks DW, Nattrass S, Cant MA, Weiss MN, Giles D, Balcomb KC, Croft DP (2017). Mortality risk and social network position in resident killer whales: Sex differences and the importance of resource abundance. Proceedings of the Royal Society B: Biological Sciences, 284(1865). Abstract.
Bolt SL, Boyland NK, Mlynski DT, James R, Croft DP (2017). Pair housing of dairy calves and age at pairing: Effects on weaning stress, health, production and social networks. PLoS ONE, 12(1). Abstract.
Croft DP, Johnstone RA, Ellis S, Nattrass S, Franks DW, Brent LJN, Mazzi S, Balcomb KC, Ford JKB, Cant MA, et al (2017). Reproductive Conflict and the Evolution of Menopause in Killer Whales. Current Biology, 27(2), 298-304. Abstract.
Silk MJ, Weber N, Steward LC, Delahay RJ, Croft DP, Hodgson DJ, Boots M, McDonald RA (2017). Seasonal variation in daily patterns of social contacts in the European badger meles meles. Ecology and Evolution, 7(21), 9006-9015. Abstract.
Rose PE, Croft DP (2017). Social bonds in a flock bird: Species differences and seasonality in social structure in captive flamingo flocks over a 12-month period. Applied Animal Behaviour Science, 193, 87-97. Abstract.
Stevens CH, Croft DP, Paull GC, Tyler CR (2017). Stress and welfare in ornamental fishes: what can be learned from aquaculture?. Journal of Fish Biology, 91(2), 409-428. Abstract.
Silk MJ, Croft DP, Delahay RJ, Hodgson DJ, Weber N, Boots M, McDonald RA (2017). The application of statistical network models in disease research. Methods in Ecology and Evolution, 8(9), 1026-1041. Abstract.
Edenbrow M, Bleakley BH, Darden SK, Tyler CR, Ramnarine IW, Croft DP (2017). The evolution of cooperation: Interacting phenotypes among social partners. American Naturalist, 189(6), 630-643. Abstract.
Silk MJ, Croft DP, Delahay RJ, Hodgson DJ, Boots M, Weber N, McDONALD RA (2017). Using social network measures in wildlife disease ecology, epidemiology, and management. BioScience, 67(3), 245-257. Abstract.
Firth JA, Voelkl B, Crates RA, Aplin LM, Biro D, Croft DP, Sheldon BC (2017). Wild birds respond to flockmate loss by increasing their social network associations to others. Proceedings of the Royal Society B: Biological Sciences, 284(1854). Abstract.
Croft DP, Darden SK, Wey TW (2016). Current directions in animal social networks. Current Opinion in Behavioral Sciences, 12, 52-58. Abstract.
Killen SS, Croft DP, Salin K, Darden SK (2016). Male sexually coercive behaviour drives increased swimming efficiency in female guppies. Functional Ecology, 30(4), 576-583. Abstract.
Franks DW, Nattrass S, Brent LJN, Whitehead H, Foote AD, Mazzi S, Ford JKB, Balcomb KC, Cant MA, Croft DP, et al (2016). The significance of postreproductive lifespans in killer whales: a comment on Robeck et al. Journal of Mammalogy, 97(3), 906-909. Abstract.
Boyland NK, Mlynski DT, James R, Brent LJN, Croft DP (2016). The social network structure of a dynamic group of dairy cows: from individual to group level patterns. Applied Animal Behaviour Science, 174, 1-10. Abstract.
Vázquez Diosdado JA, Barker ZE, Hodges HR, Amory JR, Croft DP, Bell NJ, Codling EA (2015). Classification of behaviour in housed dairy cows using an accelerometer-based activity monitoring system. Animal Biotelemetry, 3(1). Abstract.
Brent LJN, Franks DW, Foster EA, Balcomb KC, Cant MA, Croft DP (2015). Ecological knowledge, leadership, and the evolution of menopause in killer whales. Current Biology, 25(6), 746-750.
Rose PE, Croft DP (2015). Evidence of directed interactions between individuals in captive flamingo flocks. Wildfowl, 65, 121-132. Abstract.
Silk MJ, Jackson AL, Croft DP, Colhoun K, Bearhop S (2015). The consequences of unidentifiable individuals for the analysis of an animal social network. Animal Behaviour, 104, 1-11. Abstract.
Croft DP, Brent LJN, Franks DW, Cant MA (2015). The evolution of prolonged life after reproduction. TRENDS IN ECOLOGY & EVOLUTION, 30(7), 407-416.  Author URL.
Rose PE, Croft DP (2015). The potential of Social Network Analysis as a tool for the management of zoo animals. ANIMAL WELFARE, 24(2), 123-138.  Author URL.
Tranmer M, Marcum CS, Morton FB, Croft DP, de Kort SR (2015). Using the relational event model (REM) to investigate the temporal dynamics of animal social networks. Animal Behaviour, 101, 99-105. Abstract.
Rose PE, Croft DP, Lee R (2014). A review of captive flamingo (Phoenicopteridae) welfare: a synthesis of current knowledge and future directions. International Zoo Yearbook, 48(1), 139-155. Abstract.
Wilson ADM, Krause S, James R, Croft DP, Ramnarine IW, Borner KK, Clement RJG, Krause J (2014). Dynamic social networks in guppies (<i>Poecilia reticulata</i>). BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY, 68(6), 915-925.  Author URL.
Jacoby DMP, Fear LN, Sims DW, Croft DP (2014). Shark personalities? Repeatability of social network traits in a widely distributed predatory fish. BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY, 68(12), 1995-2003.  Author URL.
Kurvers RHJM, Krause J, Croft DP, Wilson ADM, Wolf M (2014). The evolutionary and ecological consequences of animal social networks: emerging issues. Trends Ecol Evol, 29(6), 326-335. Abstract.  Author URL.
Silk MJ, Croft DP, Tregenza T, Bearhop S (2014). The importance of fission-fusion social group dynamics in birds. IBIS, 156(4), 701-715.  Author URL.
Wilson ADM, Croft DP, Krause J (2013). Social networks in elasmobranchs and teleost fishes. Fish and Fisheries
Boyland NK, James R, Mlynski DT, Madden JR, Croft DP (2013). Spatial proximity loggers for recording animal social networks: Consequences of inter-logger variation in performance. Behavioral Ecology and Sociobiology, 67(11), 1877-1890. Abstract.
Foster EA, Franks DW, Mazzi S, Darden SK, Balcomb KC, Ford JKB, Croft DP (2012). Adaptive prolonged postreproductive life span in killer whales. Science, 337(6100). Abstract.  Author URL.
Keedwell E, Morley M, Croft D (2012). Continuous Trait-Based Particle Swarm Optimisation (CTB-PSO). Lecture Notes in Computer Science (including subseries Lecture Notes in Artificial Intelligence and Lecture Notes in Bioinformatics), 7461 LNCS, 342-343. Abstract.
Jacoby DMP, Brooks EJ, Croft DP, Sims DW (2012). Developing a deeper understanding of animal movements and spatial dynamics through novel application of network analyses. METHODS IN ECOLOGY AND EVOLUTION, 3(3), 574-583.  Author URL.
Bode NWF, Franks DW, Jamie Wood A, Piercy JJB, Croft DP, Codling EA (2012). Distinguishing social from nonsocial navigation in moving animal groups. American Naturalist, 179(5), 621-632. Abstract.
Edenbrow M, Croft DP (2012). Environmental and genetic effects shape the development of personality traits in the mangrove killifish Kryptolebias marmoratus. Oikos
Edenbrow M, Croft DP (2012). Kin and familiarity influence association preferences and aggression in the mangrove killifish Kryptolebias marmoratus. J Fish Biol, 80(3), 503-518. Abstract.  Author URL.
Edenbrow M, Croft DP (2012). Sequential hermaphroditism and personality in a clonal vertebrate: the mangrove killifish. Behav Processes, 90(2), 229-237. Abstract.  Author URL.
Jacoby DMP, Croft DP, Sims DW (2012). Social behaviour in sharks and rays: Analysis, patterns and implications for conservation. Fish and Fisheries, 13(4), 399-417.
Foster EA, Franks DW, Morrell LJ, Balcomb KC, Parsons KM, van Ginneken A, Croft DP (2012). Social network correlates of food availability in an endangered population of killer whales, <i>Orcinus orca</i>. ANIMAL BEHAVIOUR, 83(3), 731-736.  Author URL.
Jacoby DMP, Sims DW, Croft DP (2012). The effect of familiarity on aggregation and social behaviour in juvenile small spotted catsharks Scyliorhinus canicula. J Fish Biol, 81(5), 1596-1610. Abstract.  Author URL.
Griffiths AM, Jacoby DMP, Casane D, McHugh M, Croft DP, Genner MJ, Sims DW (2012). The first analysis of multiple paternity in an oviparous shark species, the small-spotted catshark (Scyliorhinus canicula L.). Journal of Heredity, 2(103), 166-73.
Croft DP, Hamilton PB, Darden SK, Jacoby DMP, James R, Bettaney EM, Tyler CR (2012). The role of relatedness in structuring the social network of a wild guppy population. Oecologia, 1-9.
Guttridge TL, Gruber SH, DiBattista JD, Feldheim KA, Krause S, Croft DP, Krause J (2011). Assortative interactions and leadership in free-ranging population of juvenile lemon sharks, Negaprion brevirostris. Marine Ecology Progress Series, 423, 235-245.
Edenbrow M, Croft DP (2011). Behavioural types and life history strategies during ontogeny in the mangrove killifish, Kryptolebias marmoratus. Animal Behaviour, 82, 731-741. Abstract.
Croft DP, Edenbrow M, Darden SK, Ramnarine I, van Oosterhout C, Cable J (2011). Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies Poecilia reticulata. Behavioral Ecology and Sociobiology, 65, 2219-2227. Abstract.
Mourabit S, Edenbrow M, Croft DP, Kudoh1 T (2011). Embryonic Development of the Self-fertilizing Mangrove Killifish Kryptolebias marmoratus. Developmental Dynamics(240), 1694-1704.
Edenbrow M, Darden SK, Ramnarine IW, Evans JP, James R, Croft DP (2011). Environmental effects on social interaction networks and male reproductive behaviour in guppies, Poecilia reticulata. Animal Behaviour, 81(3), 551-558. Abstract.
Croft DP, Madden J, Franks D, James R (2011). Hypothesis testing in animal social networks. Trends in Ecology and Evolution, 26, 502-507.
Krause J, Wilson AMD, Croft DP (2011). New technology facilitates the study of social networks. Trends in Ecology and Evolution, 26, 5-6.
Krause J, James R, Croft DP (2011). Personality in the context of social networks. Philosophical Transactions of the Royal Society B: Biological Sciences, 4099-4106. Abstract.
Kelley JL, Morrell LJ, Inskip C, Krause J, Croft DP (2011). Predation risk shapes social networks in fission-fusion populations. PLoS-One, 6(8)
Brask JB, Croft DP, Thompson K, Dabelsteen T, Darden S (2011). Social preferences based on sexual attractiveness: a female strategy to reduce male sexual attention. Proc Biol Sci, 279, 1748-1753. Abstract.
Piyapong C, Krause J, Chapman BB, Ramnarine IW, Louca V, Croft DP (2010). Sex matters: a social context to boldness in guppies (Poecilia reticulata). Behavioral Ecology, 21(1), 3-8. Abstract.
Jones KA, Croft DP, Ramnarine IW, Godin JGJ (2010). Size-assortative shoaling in the guppy (Poecilia reticulata): the role of active choice. Ethology, 116(2), 147-154. Abstract.
Croft DP, Krause J, Darden SK, Ramnarine IW, Faria JJ, James R (2009). Behavioural trait assortment in a social network: Patterns and implications. Behavioral Ecology and Sociobiology, 63(10), 1495-1503. Abstract.
Shorrocks B, Croft DP (2009). Necks and networks: a preliminary study of population structure in the reticulated giraffe (Giraffa camelopardalis reticulata de Winston). African Journal of Ecology, 47(3), 374-381. Abstract.
James R, Croft DP, Krause J (2009). Potential banana skins in animal social network analysis. Behavioral Ecology and Sociobiology, 63(7), 989-997. Abstract.
Croft DP, Darden SK, Ruxton GD (2009). Predation risk as a driving force for phenotypic assortment: a cross-population comparison. Proc Biol Sci, 276(1663), 1899-1904. Abstract.  Author URL.
Dyer JRG, Croft DP, Morrell LJ, Krause J (2009). Shoal composition determines foraging success in the guppy. Behavioral Ecology, 20(1), 165-171. Abstract.
Darden SK, James R, Ramnarine IW, Croft DP (2009). Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition. Proc Biol Sci, 276(1667), 2651-2656. Abstract.  Author URL.
Guttridge TL, Gruber SH, Gledhill KS, Croft DP, Sims DW, Krause J (2009). Social preferences of juvenile lemon sharks, Negaprion brevirostris. Animal Behaviour, 78(2), 543-548. Abstract.
Morrell LJ, Croft DP, Dyer JRG, Chapman BB, Kelley JL, Laland KN, Krause J (2008). Association patterns and foraging behaviour in natural and artificial guppy shoals. Animal Behaviour, 76(3), 855-864. Abstract.
Dyer JRG, Ioannou CC, Morrell LJ, Croft DP, Couzin ID, Waters DA, Krause J (2008). Consensus decision making in human crowds. Animal Behaviour, 75(2), 461-470. Abstract.
Thomas POR, Croft DP, Morrell LJ, Davis A, Faria JJ, Dyer JRG, Piyapong C, Ramnarine I, Ruxton GD, Krause J, et al (2008). Does defection during predator inspection affect social structure in wild shoals of guppies?. Animal Behaviour, 75(1), 43-53. Abstract.
Darden SK, Croft DP (2008). Male harassment drives females to alter habitat use and leads to segregation of the sexes. Biol Lett, 4(5), 449-451. Abstract.  Author URL.
Botham MS, Hayward RK, Morrell LJ, Croft DP, Ward JR, Ramnarine I, Krause J (2008). Risk-sensitive antipredator behavior in the Trinidadian guppy, Poecilia reticulata. Ecology, 89(11), 3174-3185. Abstract.
Piyapong C, Morrell LJ, Croft DP, Dyer JRG, Ioannou CC, Krause J (2007). A cost of leadership in human groups. Ethology, 113(9), 821-824. Abstract.
Morrell LJ, Hunt KL, Croft DP, Krause J (2007). Diet, familiarity and shoaling decisions in guppies. Animal Behaviour, 74(2), 311-319. Abstract.
Krause J, Croft DP, James R (2007). Social network theory in the behavioural sciences: Potential applications. Behavioral Ecology and Sociobiology, 62(1), 15-27. Abstract.
Croft DP, Morrell LJ, Wade AS, Piyapong C, Ioannou CC, Dyer JRG, Chapman BB, Wong Y, Krause J (2006). Predation risk as a driving force for sexual segregation: a cross-population comparison. Am Nat, 167(6), 867-878. Abstract.  Author URL.
Wright D, Ward AJW, Croft DP, Krause J (2006). Social organization, grouping, and domestication in fish. Zebrafish, 3(2), 141-155. Abstract.
Croft DP, James R, Thomas POR, Hathaway C, Mawdsley D, Laland KN, Krause J (2006). Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). Behavioral Ecology and Sociobiology, 59(5), 644-650. Abstract.
Croft DP, James R, Ward AJW, Botham MS, Mawdsley D, Krause J (2005). Assortative interactions and social networks in fish. Oecologia, 143(2), 211-219. Abstract.  Author URL.
Croft DP, Arrowsmith BJ, Webster M, Krause J (2004). Intra-sexual preferences for familiar fish in male guppies. Journal of Fish Biology, 64(1), 279-283. Abstract.
Croft DP, Botham MS, Krause J (2004). Is sexual segregation in the guppy, Poecilia reticulata, consistent with the predation risk hypothesis?. Environmental Biology of Fishes, 71(2), 127-133. Abstract.
Croft DP, Krause J, James R (2004). Social networks in the guppy (Poecilia reticulata). Proc Biol Sci, 271 Suppl 6(Suppl 6), S516-S519. Abstract.  Author URL.
Croft DP, Arrowsmith BJ, Bielby J, Skinner K, White E, Couzin ID, Magurran AE, Ramnarine I, Krause J (2003). Mechanisms underlying shoal composition in the Trinidadian guppy, Poecilia reticulata. Oikos, 100(3), 429-438. Abstract.
Croft DP, Albanese B, Arrowsmith BJ, Botham M, Webster M, Krause J (2003). Sex-biased movement in the guppy (Poecilia reticulata). Oecologia, 137(1), 62-68. Abstract.  Author URL.
Croft DP, Krause J, Couzin ID, Pitcher TJ (2003). When fish shoals meet: Outcomes for evolution and fisheries. Fish and Fisheries, 4(2), 138-146. Abstract.
Hoare DJ, Ward AJW, Couzin ID, Croft DP, Krause J (2001). A grid-net technique for the analysis of fish positions within free-ranging shoals. Journal of Fish Biology, 59(6), 1667-1672. Abstract.
Krause J, Hoare DJ, Croft D, Lawrence J, Ward A, Ruxton GD, Godin JG, James R (2000). Fish shoal composition: mechanisms and constraints. Proc Biol Sci, 267(1456), 2011-2017. Abstract.  Author URL.


Pavelka MSM, Brent LJN, Croft DP, Fedigan LM (2018). Post-fertile Lifespan in Female Primates and Cetaceans. In Kalbitzer U, Jack K (Eds.) Primate Life Histories, Sex Roles, and Adaptability. Developments in Primatology: Progress and Prospects, Springer, Cham.
Croft DP, Edenbrow M, Darden SK (2014). Assortment in social networks and the evolution of cooperation. In Krause J, James R, Franks D, Croft DP (Eds.) Animal Social Networks, Oxford University Press.
Ioannou CC, Couzin ID, James R, Croft DP, Krause J (2011). Social Organisation and Information Transfer in Schooling Fish. In  (Ed) Fish Cognition and Behavior, 217-239.
Krause J, Ward AJW, James R, Croft DP (2008). Group living and social networks. In C.Magnhagen, V.Braithwaite, E.Forsgren, Kapoor & BG (eds). In Magnhagen C, Braithwaite V, Forsgren E, Kapoor BG (Eds.) Fish behaviour.
Krause J, Ward AJW, James R, Croft DP (2008). Group-living and social networks. In  (Ed) Fish Behaviour, 485-498. Abstract.
Croft DP, Griffiths SW, Magurran AE, Krause J (2007). The Influence of Reproductive Behaviour on the Social Organisation of Shoaling Fish. In Rocha MJ, Arukwe A, Kapoor BG (Eds.) Fish Reproduction.
Krause J, James R, Croft DP (2006). Analyse sozialer Netzwerke. In  (Ed) Methoden der Verhaltensbiologie, Springer Nature, 148-152.
Croft DP, Krause J, James R (2006). Predator avoidance and reproductive strategies: Predation risk as a driving factor for size assortative shoaling and its implications for sexual segregation in fish. In  (Ed) Sexual Segregation in Vertebrates: Ecology of the Two Sexes, 115-126. Abstract.
Couzin ID, James R, Croft DP, Krause J (2006). Social organization and information transfer in schooling fish. In Brown C, Laland KN, Krause J (Eds.) Fish Cognition and Behaviour, Cambridge: Blackwell Publishing.
Krause J, James R, Croft DP (2006). Soziale Netzwerke bei Tieren. In Naguib M (Ed) Methoden der Verhaltensbiologie Heidelberg, Springer-Verlag.
Croft DP, James R, Krause J (2005). Predator avoidance as a factor for size assortative shoaling and its implications for sex segregation in fish. In Ruckstuhl K, Neuhaus P (Eds.) Sexual segregation in vertebrates: ecology of the two sexes, Cambridge University Press.

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External Engagement and Impact


2014: Fisheries of the British Isles Research Medal: awarded to younger scientists in recognition of exceptional advances in the study of fish biology and/or fisheries.

Editorial responsibilities

Associate Editor, Behavioral Ecology & Sociobiology (2011-).

Ethics committee for Animal Behaviour (2012-2017). 

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PSY3401 Psychology Research Project (BSc)

PSYM210 Research Apprehenticeship (MSc)

PSY2217 Wild Behaviour (15 credits)



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Supervision / Group

Postdoctoral researchers

Postgraduate researchers


  • Jade Barrett
  • Natasha Boyland
  • Destiny Bradley
  • Lauren Brent
  • Mathew Edenbrow PhD 2011
  • Emma Foster PhD 2012
  • Robert Heathcote
  • David Jacoby PhD 2012
  • Alessandro Macario PhD 2013
  • Paul Rose
  • Chloe Stevens
  • Dawn Thomas PhD 2011
  • Jon Ward PhD 2010

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Office Hours:

Monday 11:00-12:00am

Tuesday 11:30-12:30 am 

Book chat during office hours

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