Professor Edward Codling

Faculty Dean Research (Science and Health) - Professor (R)
Department of Mathematical Sciences
Professor Edward Codling
  • Email

  • Telephone

    +44 (0) 1206 874567

  • Location

    6.315, Colchester Campus

  • Academic support hours

    Open door policy



I joined the Department of Mathematical Sciences at the University of Essex in 2006, and I'm currently a Professor of Mathematical Biology. From August 2020 I will be undertaking the role of Faculty Dean for Research in the Faculty of Science and Health. My research is focused on using novel mathematical and computational techniques to study problems in biology, ecology, and animal and environmental sciences. In particular, my main interest is in movement ecology, and specifically the development of theoretical models and empirical analysis tools that give insights into animal (including human) movement and behaviour at both the individual and collective level. My current main focus is on developing and using novel tracking technology and modelling tools to analyse the movement and behaviour of managed animals (dairy cows in particular) in order to gain insights into health, welfare and to improve management. I am also interested in how movement influences spatial population dynamics, and the application of modelling and analysis tools to other natural resource management questions (e.g. in fisheries). The truly inter-disciplinary nature of my research means I work with a range of collaborators from different fields and backgrounds. I passionately believe in the value of communicating scientific research, and the role that mathematics and quantitative approaches can play in such research, across disciplines and to the wider public. To this end, I have organised and participated in a wide range of training workshops, high profile public engagement events and media activities. I have also worked closely with public sector and industry on a number of practical applications of my research. I am an elected Fellow of the Institute of Mathematics and its Applications (FIMA) and an elected Fellow of the Royal Society of Biology (FRSB). I am also a Senior Fellow of the Higher Education Academy (SFHEA).


  • PhD University of Leeds, (2003)


University of Essex

  • Faculty Dean for Research, Faculty of Science and Health, University of Essex (8/2020 - present)

  • Professor of Mathematical Biology, Mathematical Sciences, University of Essex (1/10/2014 - present)

  • Senior Lecturer in Mathematical Biology, Mathematical Sciences, University of Essex (1/10/2011 - 1/9/2014)

  • Lecturer in Mathematical Biology, Mathematical Sciences, University of Essex (1/9/2006 - 1/9/2011)

Other academic

  • Postdoctoral Fellow, University College Cork & Marine Institute (Galway) (2/2004 - 8/2006)

Research and professional activities

Research interests

Animal movement and dispersal

Open to supervise

Random walks and diffusion

Open to supervise

Analysis of movement data

Open to supervise

Collective behaviour

Open to supervise

Human crowd behaviour

Open to supervise

Animal social network analysis

Open to supervise

Fisheries modelling, assessment, and management

Open to supervise

Modelling population dynamics

Open to supervise

Current research

Movement and Behavioural Ecology: modelling the behaviour, movement and dispersal of animals, micro-organisms and cells.

Population Ecology: mathematical analysis and simulation of the population dynamics and optimal management strategies of fisheries and marine ecosystems.

Teaching and supervision

Previous supervision

Joseph David Bailey
Joseph David Bailey
Thesis title: Modelling and Analysis of Individual Animal Movement
Degree subject: Mathematics
Degree type: Doctor of Philosophy
Awarded date: 9/1/2020
Tahani Ali Salman Al-Karkhi
Tahani Ali Salman Al-Karkhi
Thesis title: Population Dynamics and Pattern Formation in an Info-Chemical Mediated Tri-Trophic Plankton Model
Degree subject: Applied Mathematics
Degree type: Doctor of Philosophy
Awarded date: 15/8/2018
Alexander Kieran Partner
Alexander Kieran Partner
Thesis title: A Hybrid Cusum Approach to Identify Residency and Transition Periods for Animal Movement with an Application to Housed Dairy Cows
Degree subject: Applied Mathematics
Degree type: Master of Science (by Dissertation)
Awarded date: 14/5/2018


Journal articles (58)

Bailey, JD., Benefer, CM., Blackshaw, RP. and Codling, EA., (2021). Walking behaviour in the ground beetle, Poecilus cupreus: dispersal potential, intermittency and individual variation. Bulletin of Entomological Research. 111 (2), 200-209

Bailey, JD. and Codling, EA., (2021). Emergence of the wrapped Cauchy distribution in mixed directional data. AStA Advances in Statistical Analysis. 105 (2), 229-246

Bailey, JD., King, AJ., Codling, EA., Short, AM., Johns, GI. and Fürtbauer, I., (2021). “Micropersonality” traits and their implications for behavioral and movement ecology research. Ecology and Evolution. 11 (7), 3264-3273

Miller, GA., Mitchell, M., Barker, ZE., Giebel, K., Codling, EA., Amory, JR., Michie, C., Davison, C., Tachtatzis, C., Andonovic, I. and Duthie, C-A., (2020). Using animal-mounted sensor technology and machine learning to predict time-to-calving in beef and dairy cows. animal. 14 (6), 1304-1312

Al-Karkhi, TAS., Kusdiantara, R., Susanto, H. and Codling, EA., (2020). Bloom Formation and Turing Patterns in an Infochemical Mediated Multitrophic Plankton Model. International Journal of Bifurcation and Chaos. 30 (10), 2030028-2030028

Chopra, K., Hodges, HR., Barker, ZE., Vázquez Diosdado, JA., Amory, JR., Cameron, TC., Croft, DP., Bell, NJ. and Codling, EA., (2020). Proximity Interactions in a Permanently Housed Dairy Herd: Network Structure, Consistency, and Individual Differences. Frontiers in Veterinary Science. 7, 583715-

Bode, NWF. and Codling, EA., (2019). Exploring Determinants of Pre-movement Delays in a Virtual Crowd Evacuation Experiment. Fire Technology. 55 (2), 595-615

Watson, FM., Hepburn, LJ., Cameron, T., Le Quesne, WJF. and Codling, EA., (2019). Relative mobility determines the efficacy of MPAs in a two species mixed fishery with conflicting management objectives. Fisheries Research. 219, 105334-105334

Walker, N., Susanto, H., Steinke, M. and Codling, EA., (2019). Bottom-up and top-down control in a multitrophic system: the role of nutrient limitation and infochemical-mediated predation in a plankton food-web model. Communication in Biomathematical Sciences. 2 (2), 65-84

Bailey, JD., Wallis, J. and Codling, EA., (2018). Navigational efficiency in a biased and correlated random walk model of individual animal movement. Ecology. 99 (1), 217-223

Berdahl, AM., Kao, AB., Flack, A., Westley, PAH., Codling, EA., Couzin, ID., Dell, AI. and Biro, D., (2018). Collective animal navigation and migratory culture: From theoretical models to empirical evidence. Philosophical Transactions of the Royal Society B: Biological Sciences. 373 (1746), 20170009-20170009

Piercy, JJB., Smith, DJ., Jompa, J., Simpson, SD. and Codling, EA., (2018). High temporal resolution sampling reveals reef fish settlement is highly clustered. Canadian Journal of Fisheries and Aquatic Sciences. 75 (4), 560-568

Barker, ZE., Vázquez Diosdado, JA., Codling, EA., Bell, NJ., Hodges, HR., Croft, DP. and 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

Vázquez Diosdado, JA., Barker, ZE., Hodges, HR., Amory, JR., Croft, DP., Bell, NJ. and 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), e0208424-e0208424

Piercy, JJB., Smith, DJ., Codling, EA., Hill, AJ. and Simpson, SD., (2016). The Good, The Bad, and The Distant: Soundscape Cues for Larval Fish. EFFECTS OF NOISE ON AQUATIC LIFE II. 875, 829-837

Auger-Méthé, M., Derocher, AE., DeMars, CA., Plank, MJ., Codling, EA. and Lewis, MA., (2016). Evaluating random search strategies in three mammals from distinct feeding guilds. Journal of Animal Ecology. 85 (5), 1411-1421

Pazhayamadom, DG., Kelly, CJ., Rogan, E. and Codling, EA., (2016). Self-starting cumulative sum harvest control rule (SS-CUSUM-HCR) for status-quo management of data-limited fisheries. Canadian Journal of Fisheries and Aquatic Sciences. 73 (3), 366-381

Codling, EA. and Bode, NWF., (2016). Balancing direct and indirect sources of navigational information in a leaderless model of collective animal movement. Journal of Theoretical Biology. 394, 32-42

Auger-Méthé, M., Derocher, AE., Plank, MJ., Codling, EA. and Lewis, MA., (2015). Differentiating the Lévy walk from a composite correlated random walk. Methods in Ecology and Evolution. 6 (10), 1179-1189

Bode, NWF., Miller, J., O'Gorman, R. and Codling, EA., (2015). Increased costs reduce reciprocal helping behaviour of humans in a virtual evacuation experiment. Scientific Reports. 5 (1), 15896-

Pazhayamadom, DG., Kelly, CJ., Rogan, E. and Codling, EA., (2015). Decision Interval Cumulative Sum Harvest Control Rules (DI-CUSUM-HCR) for managing fisheries with limited historical information. Fisheries Research. 171, 154-169

Kraak, SBM., Reid, DG., Bal, G., Barkai, A., Codling, EA., Kelly, CJ. and Rogan, E., (2015). RTI (“Real-Time Incentives”) outperforms traditional management in a simulated mixed fishery and cases incorporating protection of vulnerable species and areas. Fisheries Research. 172, 209-224

Vázquez Diosdado, JA., Barker, ZE., Hodges, HR., Amory, JR., Croft, DP., Bell, NJ. and Codling, EA., (2015). Classification of behaviour in housed dairy cows using an accelerometer-based activity monitoring system. Animal Biotelemetry. 3 (1)

Bode, NWF., Kemloh Wagoum, AU. and Codling, EA., (2015). Information use by humans during dynamic route choice in virtual crowd evacuations. Royal Society Open Science. 2 (1), 140410-140410

Piercy, JJB., Codling, EA., Hill, AJ., Smith, DJ. and Simpson, SD., (2014). Habitat quality affects sound production and likely distance of detection on coral reefs. Marine Ecology Progress Series. 516, 35-47

Codling, EA., (2014). Pest insect movement and dispersal as an example of applied movement ecology. Physics of Life Reviews. 11 (3), 533-535

Kraak, SBM., Reid, DG. and Codling, EA., (2014). Exploring the RTI (real-time incentive) tariff-based approach to single-species fisheries management. Fisheries Research. 155, 90-102

Auger-Méthé, M., Plank, MJ. and Codling, EA., (2014). Distinguishing between Lévy walks and strong alternative models: comment. Ecology. 95 (4), 1104-1109

Codling, EA. and Bode, NWF., (2014). Copycat dynamics in leaderless animal group navigation. Movement Ecology. 2 (1), creators-Codling=3AEdward_A=3A=3A

Bode, NWF., Kemloh Wagoum, AU. and Codling, EA., (2014). Human responses to multiple sources of directional information in virtual crowd evacuations. Journal of the Royal Society Interface. 11 (91), 20130904-20130904

Breckels, MN., Bode, NWF., Codling, EA. and Steinke, M., (2013). Effect of grazing-mediated dimethyl sulfide (DMS) production on the swimming behavior of the copepod Calanus helgolandicus.. Marine Drugs. 11 (7), 2486-2500

Lewis, ND., Morozov, A., Breckels, MN., Steinke, M. and Codling, EA., (2013). Multitrophic Interactions in the Sea: Assessing the Effect of Infochemical-Mediated Foraging in a 1-d Spatial Model. Mathematical Modelling of Natural Phenomena. 8 (6), 25-44

Lewis, ND., Breckels, MN., Steinke, M. and Codling, EA., (2013). Role of infochemical mediated zooplankton grazing in a phytoplankton competition model. Ecological Complexity. 16, 41-50

Codling, EA. and Dumbrell, AJ., (2013). Bridging the gap between theory and data in ecological models. Ecological Complexity. 16, 1-8

Simpson, SD., Piercy, JJB., King, J. and Codling, EA., (2013). Modelling larval dispersal and behaviour of coral reef fishes. Ecological Complexity. 16, 68-76

Bode, NWF. and Codling, EA., (2013). Human exit route choice in virtual crowd evacuations. Animal Behaviour. 86 (2), 347-358

Pazhayamadom, DG., Kelly, CJ., Rogan, E. and Codling, EA., (2013). Self-starting CUSUM approach for monitoring data poor fisheries. Fisheries Research. 145, 114-127

Lewis, ND., Breckels, MN., Archer, SD., Morozov, A., Pitchford, JW., Steinke, M. and Codling, EA., (2012). Grazing-induced production of DMS can stabilize food-web dynamics and promote the formation of phytoplankton blooms in a multitrophic plankton model. Biogeochemistry. 110 (1-3), 303-313

Codling, EA. and Dumbrell, AJ., (2012). Mathematical and theoretical ecology: linking models with ecological processes. Interface Focus. 2 (2), 144-149

Bode, NWF., Franks, DW., Wood, AJ., Piercy, JJB., Croft, DP. and Codling, EA., (2012). Distinguishing Social from Nonsocial Navigation in Moving Animal Groups. The American Naturalist. 179 (5), 621-632

Knell, AS. and Codling, EA., (2012). Classifying area-restricted search (ARS) using a partial sum approach. Theoretical Ecology. 5 (3), 325-339

Kraak, SBM., Reid, DG., Gerritsen, HD., Kelly, CJ., Fitzpatrick, M., Codling, EA. and Rogan, E., (2012). 21st century fisheries management: a spatio-temporally explicit tariff-based approach combining multiple drivers and incentivising responsible fishing. ICES Journal of Marine Science. 69 (4), 590-601

Boakes, DE., Codling, EA., Thorn, GJ. and Steinke, M., (2011). Analysis and modelling of swimming behaviour in Oxyrrhis marina. Journal of Plankton Research. 33 (4), 641-649

Plank, MJ. and Codling, EA., (2011). Sampling rate and misidentification of Lévy and non-Lévy movement paths: reply. Ecology. 92 (8), 1701-1702

Codling, EA. and Plank, MJ., (2011). Turn designation, sampling rate and the misidentification of power laws in movement path data using maximum likelihood estimates. Theoretical Ecology. 4 (3), 397-406

Breckels, MN., Boakes, DE., Codling, EA., Malin, G., Archer, SD. and Steinke, M., (2010). Modelling the concentration of exuded dimethylsulphoniopropionate (DMSP) in the boundary layer surrounding phytoplankton cells. Journal of Plankton Research. 32 (2), 253-257

Codling, EA., Bearon, RN. and Thorn, GJ., (2010). Diffusion about the mean drift location in a biased random walk. Ecology. 91 (10), 3106-3113

Kraak, SBM., Kelly, CJ., Codling, EA. and Rogan, E., (2010). On scientists’ discomfort in fisheries advisory science: the example of simulation-based fisheries management-strategy evaluations. Fish and Fisheries. 11 (2), 119-132

Plank, MJ. and Codling, EA., (2009). Sampling rate and misidentification of Lévy and non-Lévy movement paths. Ecology. 90 (12), 3546-3553

Faria, JJ., Codling, EA., Dyer, JRG., Trillmich, F. and Krause, J., (2009). Navigation in human crowds; testing the many-wrongs principle. Animal Behaviour. 78 (3), 587-591

Codling, EA., Plank, MJ. and Benhamou, S., (2008). Random walk models in biology. Journal of The Royal Society Interface. 5 (25), 813-834

Codling, EA., Pitchford, JW. and Simpson, SD., (2007). GROUP NAVIGATION AND THE “MANY-WRONGS PRINCIPLE” IN MODELS OF ANIMAL MOVEMENT. Ecology. 88 (7), 1864-1870

Pitchford, JW., Codling, EA. and Psarra, D., (2007). Uncertainty and sustainability in fisheries and the benefit of marine protected areas. Ecological Modelling. 207 (2-4), 286-292

Kelly, CJ., Codling, EA. and Rogan, E., (2006). The Irish Sea cod recovery plan: some lessons learned. ICES Journal of Marine Science. 63 (4), 600-610

Kelly, CJ. and Codling, EA., (2006). ‘Cheap and dirty’ fisheries science and management in the North Atlantic. Fisheries Research. 79 (3), 233-238

Codling, EA. and Hill, NA., (2005). Sampling rate effects on measurements of correlated and biased random walks. Journal of Theoretical Biology. 233 (4), 573-588

Codling, EA. and Hill, NA., (2005). Calculating spatial statistics for velocity jump processes with experimentally observed reorientation parameters. Journal of Mathematical Biology. 51 (5), 527-556

Codling, EA., Hill, NA., Pitchford, JW. and Simpson, SD., (2004). Random walk models for the movement and recruitment of reef fish larvae. Marine Ecology Progress Series. 279, 215-224

Book chapters (1)

Plank, MJ., Auger-Méthé, M. and Codling, EA., (2013). Lévy or Not? Analysing Positional Data from Animal Movement Paths. In: Dispersal, Individual Movement and Spatial Ecology. Editors: Lewis, MA., Maini, PK. and Petrovskii, SV., . Springer. 33- 52. 9783642354960

Thesis dissertation (1)

Codling, EA., (2010). Biased random walks in biology.

Other (1)

Bode, NWF., Franks, DW., Wood, AJ., Piercy, JJB., Croft, DP. and Codling, EA., (2012).Data from: Distinguishing social from nonsocial navigation in moving animal groups, American Naturalist 179(5): 621-632. doi:10.1086/665005,Dryad Digital Repository

Grants and funding


Support of collaborative research with Prof Michael Plank at The University of Essex, Colchester.

London Mathematical Society


Novel animal-mounted sensor technology to improve efficiency and sustainability

Natural Environment Research Council


Individual Specific Risks in Evacuation: Women and Children first?

AXA Research Fund

Assessment of Dairy Cow Welfare through Predictive Modelling of Individual and Social Behaviour

Biotechnology & Biology Science Res.Council


Understanding Mathematical Models in Ecology

British Ecological Society (BES)


Role of Dimethyl Sulphide

Natural Environment Research Council


Simplifying Fisheries & Natural Resources

Natural Environment Research Council

Marine Institute research Grants - UCC

Marine Institute/University Of Cork

(DRA4500 - Finished) MI Projects via University College Cork

Marine Institute/University Of Cork

+44 (0) 1206 874567


6.315, Colchester Campus

Academic support hours:

Open door policy