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Dr Safi Darden

Dr Safi Darden

Senior Lecturer


 Washington Singer 124b


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


I work in both vertebrate and invertebrate systems and in the lab and field to investigate the causes and consequences of social behaviour. I am currently particularly interested in the evolution and maintenance of cooperation among unrelated individuals and the mechanisms underlying decision-making in cooperative contexts. 

I am part of Exeter's first cohort of WHEN's 100 Black Women Professors NOW programme so please ask me about this! If you are a student at UoE and interested in helping us with an outreach project we are running as a cohort then get in touch and also consider applying to this funded part-time research internship.

If you are looking for opportunities to come and work with me I currently have positions available - see below and on my research page and be sure to connect with me on LinkedIn or via email.

In addition to my research endeavours and teaching and supervisory activities, I am passionate about contributing to service activities that foster a more inclusive and socially just research and education environment. My service activities currently include the following roles:


PhD, Animal Behaviour, December 2006
University of Copenhagen, Institute of Biology, Copenhagen, DK

MSc, Animal Behaviour, February 2002
University of Copenhagen, Institute of Biology, Copenhagen, DK

BSc, Biology, May 1996
Cornell University, College of Agriculture and Life Sciences, Ithaca, NY, USA


Senior Lecturer, University of Exeter, Psychology, 2018-Present

Lecturer, University of Exeter, Psychology, 2013-2018

Leverhulme Research Fellow, University of Exeter, Psychology, 2011-2013

Career Break (maternity leave), 2010-2011

Teaching Fellow in Animal Behaviour, University of Exeter, Psychology, 2009-2010

Postdoctoral Research Fellow, 2009; Sexual conflict in the Trinidadian guppy; University of Exeter, School of Psychology

Career Break (maternity leave), 2008-2009

Postdoctoral Research Fellow, Sexual segregation in the Trinidadian guppy; Bangor University, School of Biological Sciences, 2007-2008


Research group links

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Research interests

In my research I investigate the interplay between the behaviour of individuals and the processes and patterns expressed at the population level. My main line of research is aimed at understanding how inter-individual interactions act as a driving force for social evolution. Using the Trinidadian guppy (Poecilia reticulata) as a model system, I have developed a novel research framework for exploring selection for social traits in the context of sexual conflict (i.e. the conflict of interest between the sexes over reproduction) and cooperation. A major component of this work lies in identifying the underlying mechanisms driving changes in social behaviour.

Research projects

I am currently taking on PhD and Masters by Research students. Any specific projects are listed below, but you can also get in touch with other project ideas. I am currently offering projects in three study systems: Trinidadian guppies (Poecilia reticulata), fiddler crabs (Afruca tangeri), and red foxes (Vulpes vulpes), but contact me if you have another system in mind.

  • Funded opportunity: PhD Studentship - 27th March 2024 deadline - Funded PhD studentship in snapping shrimp behavioural and acoustic ecology
  • Funded opportunity: PhD Scholarship - 1st April 2024 deadline - University of Exeter PhD Scholarships for Black British Researchers in the Faculty of Health and Life Sciences - get in touch to develop a project proposal and apply! Also look here for possible projects:
    • Project title: The gut microbiome and social behaviour

      A ubiquitous feature of our own evolution and that of all multicellular organisms, is that this has occurred in close contact with microbe-rich environments. For most animal species this has resulted in an existence where close associations with microorganisms feature in tissue physiology, key developmental trajectories and even expression of behaviour. Gut microbiota are thought to be the most diverse of the associated animal host microbiota and have been implicated in brain physiology and development in a diverse range of taxa, generating what has been termed a microbiota-gut-brain axis. The breadth and strength of the implications of the microbiota-gut-brain axis for behaviour are just starting to be understood, although almost exclusively in laboratory settings. From this work we know that microbes play a role in the phenotypic expression of behaviour in both vertebrate and invertebrate species and that there are feedback loops between behaviour and microbiome characteristics, including for social behaviour. Laboratory studies and literature on wild populations suggests that mechanisms of microbiome transmission and feedback along the microbiota-gut-brain axis are likely to be fundamental to a number of behavioural and ecological processes. However, the causal processes underpinning microbiome variation in wild populations and, importantly, the consequences of this variation, are currently relatively underexplored. The aim of this project is carry out empirical work on the causal and reciprocal links between sociality and gut-microbiota in free-roaming animal populations. The work can be carried out in one of a number of systems depending on the candidates interests.

      Project title: Affective states and decision making in cooperative contexts

      Across a number of species individuals monitor the behaviour of others to gauge their performance and use this information in making decisions about how to interact with their environment. Appraisal of the environment is a key component of the decision-making process. Information within the environment will vary in extent, temporal stability and temporal spacing; each with its own degree of certainty or reliability. Recent work suggests that emotions or emotion-like states are important in this process across a number of species, including humans. We might consider emotions as the culmination of both physiological and cognitive processes resulting from how individuals appraise their environment and as such, key drivers of subsequent behavioural decisions. One context where individuals monitor ‘performance’ is during cooperative interactions. Cooperation occurs when individuals perform costly behaviours that benefit one or more other individuals and is a behaviour that is ubiquitous across scales of organismal complexity (i.e. it occurs from microbes to humans). The leading hypotheses for the evolution and maintenance of cooperation among non-kin, revolve around some form of reciprocity where cooperative behaviour is rewarded at some point in the future. We propose that for reciprocity to occur, the experience of cooperative behaviour has to have a ‘positive’ social effect on receivers of cooperative behaviour or at least bystanders to a cooperative act. However, the experience of defection is likely to undermine such ‘positive’ effects that would make the proximate persistence of cooperation within a population difficult to explain. The aim of this project is to use the Trinidadian guppy (Poecilia reticulata) as a model system to investigate the affective states of individuals in response to the cooperative behaviour of others and the resultant social decisions to provide insight into the proximate psychological processes and behavioural rules that support the persistence of cooperation.

  • Funded opportunity: PhD ScholarshipUniversity of Southampton Black Futures PhD scholarship scheme - get in touch with me to develop a project with us (Principle supervisor: Dr Lauren Nadler, University of Southampton) on fish physiology and behaviour

I also regularly take on research interns - get in touch if you would like ot speak about internship opportunities. Any currently available funded opportunities are listed below.

Research grants

  • 2015 Leverhulme Trust
    The evolution of eye salience as a signal for communication
  • 2013 FNU (The Danish Council for Independent Research / Natural S
    Social niche construction and evolutionary implications for animal behaviour
  • 2010 Association for the Study of Animal Behaviour
    Research Grant: Neighbourhood effects associated with social signalling: an investigation with the European fiddler crab (Uca tangeri)
  • 2010 International Society for Behavioral Ecology
    Travel grant: Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition
  • 2010 Leverhulme Trust
    Early Career Fellowship: Social implications of the battle of the sexes
  • 2010 FNU (The Danish Council for Independent Research / Natural S
    Cognitive and physical aspects of animal communication networks

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Journal articles

Dimitriadou S, Winberg S, Thörnqvist P-O, Croft D, Darden S (In Press). Brain monoaminergic activity during predator inspection in female Trinidadian guppies (Poecilia reticulata). Behavioural Brain Research
Ciftci E, Barreto M, Doyle D, van Breen J, Darden S (In Press). Distancing or drawing together: Sexism and organizational tolerance of sexism impact women’s social relationships at work. European Journal of Social Psychology
Macario A, Darden S, Verbruggen F, Croft D (In Press). Intraspecific variation in inhibitory motor control in guppies, Poecilia reticulata. Journal of Fish Biology
Darden S-K, James R, Cave JM, Brask JB, Croft DP (In Press). Trinidadian guppies use a social heuristic that can support cooperation among non-kin. Proceedings of the Royal Society B: Biological Sciences
Padget RFB, Fawcett TW, Darden SK (2023). Guppies in large groups cooperate more frequently in an experimental test of the group size paradox. Proceedings of the Royal Society B: Biological Sciences, 290, 20230790-20230790. Abstract.
Wilde JA, Rodríguez-Muñoz R, Darden SK, Tregenza T, Fawcett TW (2023). Signalling males increase or decrease their calling effort according to the proximity of rivals in a wild cricket. Animal Behaviour, 203, 53-61. Abstract.
Dimitriadou S, Santos EM, Croft DP, van Aerle R, Ramnarine IW, Filby AL, Darden SK (2021). Social partner cooperativeness influences brain <i>oxytocin</i> transcription in Trinidadian guppies (<i>Poecilia reticulata</i>). Abstract.
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.
Darden SK, James R, Cave JM, Brask JB, Croft DP (2020). Trinidadian guppies use a social heuristic that can support cooperation among non-kin: Heuristics and real-world cooperation. Proceedings of the Royal Society B: 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. ROYAL SOCIETY OPEN SCIENCE, 6(1).  Author URL.
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.  Author URL.
Darden SK, May MK, Boyland NK, Dabelsteen T (2019). Territorial defense in a network: audiences only matter to male fiddler crabs primed for confrontation. BEHAVIORAL ECOLOGY, 30(2), 336-340.  Author URL.
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.  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.
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(1). Abstract.
Edenbrow M, Bleakley BH, Darden SK, Tyler CR, Ramnarine IW, Croft DP (2017). The Evolution of Cooperation: Interacting Phenotypes among Social Partners. The American Naturalist, 189(6), 630-643.
Croft DP, Darden SK, Wey TW (2016). Current directions in animal social networks. Current Opinion in Behavioral Sciences, 12, 52-58.
Killen SS, Croft DP, Salin K, Darden SK (2015). Male sexually coercive behaviour drives increased swimming efficiency in female guppies. Functional Ecology, 30(4), 576-583. 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.
Darden SK, Watts L (2012). Male sexual harassment alters female social behaviour towards other females. Biology Letters, 8(2), 186-188. Abstract.
Lebsock AA, Burdett CL, Darden SK, Dabelsteen T, Antolin MF, Crooks KR (2012). Space use and territoriality in swift foxes (Vulpes velox) in northeastern Colorado. Canadian Journal of Zoology, 90(3), 337-344. Abstract.
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.
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.
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.
Hebert L, Darden SK, Pedersen BV, Dabelsteen T (2011). Increased DNA amplification success of non-invasive genetic samples by successful removal of inhibitors from faecal samples collected in the field. Conservation Genetics Resources, 3(1), 41-43. Abstract.
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.
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.
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.
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.
Darden SK, Dabelsteen T (2008). Acoustic territorial signalling in a small, socially monogamous canid. Animal Behaviour, 75(3), 905-912. Abstract.
Darden SK, Steffensen LK, Dabelsteen T (2008). Information transfer among widely spaced individuals: latrines as a basis for communication networks in the swift fox?. Animal Behaviour, 75(2), 425-432. 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.
Darden SK, Pedersen SB, Larsen ON, Dabelsteen T (2008). Sound transmission at ground level in a short-grass prairie habitat and its implications for long-range communication in the swift fox Vulpes velox. J Acoust Soc Am, 124(2), 758-766. Abstract.  Author URL.
Poesel A, Dabelsteen T, Darden SK, Delhey K, Peters A (2007). Territorial responses of male blue tits, Cyanistes caeruleus, to UV-manipulated neighbours. Journal of Ornithology, 148(2), 179-187. Abstract.
Bremner-Harrison S, Harrison SWR, Cypher BL, Murdoch JD, Maldonado J, Darden SK (2006). Development of a single-sampling noninvasive hair snare. Wildlife Society Bulletin, 34(2), 456-461. Abstract.
Darden SK, Dabelsteen T (2006). Ontogeny of swift fox Vulpes velox vocalizations: Production, usage and response. Behaviour, 143(6), 659-681. Abstract.
Darden SK, Dabelsteen T, Pedersen SB (2003). A potential tool for swift fox (Vulpes velox) conservation: Individuality of long-range barking sequences. Journal of Mammalogy, 84(4), 1417-1427. Abstract.
Darden SK, Pedersen SB, Dabelsteen T (2003). Methods of frequency analysis of a complex mammalian vocalisation. Bioacoustics, 13(3), 247-263. Abstract.
Darden S, Dabelsteen T, Pedersen SB (2003). Methods of frequency analysis of a complex mammalian vocalisation. Bioacoustics 13, 247-263. Bioacoustics, 13, 247-263.


Croft DP, Edenbrow M, Darden S (2014). Assortment in social networks and the evolution of cooperation. In  (Ed) Animal Social Networks, Oxford University Press, 13-23.

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Supervision / Group

Postdoctoral researchers

Postgraduate researchers


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Office Hours:

2023/24 Term 2 Office hours: Tuesdays 14:00-16:00 - please book for an on-line or in-person slot

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