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Faculty of Health and Life Sciences

Professor Martin Stevens

Professor Martin Stevens

Professor of Sensory and Evolutionary Ecology

 01326 259358

 SERSF 1:21


Science and Engineering Research Support Facility (SERSF):, University of Exeter,  Penryn Campus, Penryn, Cornwall, TR10 9FE, UK


I am currently Director of Business, Engagement, and Innovation for the Centre for Ecology and Conservation.

My work covers sensory ecology and evolution, especially vision and adaptive coloration. Much of our current research focusses on coloration, behaviour, and human impacts in marine invertebrates.

Please visit my lab page for more information.

Recent news: my new book: Life in Colour to accompany the recent David Attenborough BBC 1 TV series is now out (see here).

In 2020 I was awarded the Zoological Society of London's Science Medal.

The research conducted in my group and with our collaborators covers a broad range of areas, including:

  • adaptive coloration and behaviour in marine animals, especially intertidal invertebrates, and human impacts on this
  • animal vision, in both the natural world and increasingly in an applied context
  • methods to analyse and quantify visual information and animal vision, especially from digital images
  • animal colour change and camouflage
  • anti-predator coloration (camouflage, warning signals, and eyespots)
  • benefits of understadning animal vision for improiving animal welfare, safety, and training.

A major current area of research is camouflage in marine species and the mechanisms and function of colour change. We are also investigating how humans are impacting coloration and behaviour through noise and chemical pollution, and invasive species. In addition, I have other applied projects, such as related to horse vision and safety in horse sports.

I work on a wide range of taxonomic groups, currently mostly marine invertebrates, but including birds, reptiles, crabs, insects, primates, and even humans. My work combines empirical and theoretical work in the lab and field, both in the UK and abroad (Africa, mainland Europe, SE Asia), and in particular locally around Cornwall. It is interdisciplinary, incorporating theories and methods from several areas of biology, experimental psychology, and computer science. Please get in touch if you are interested in joining us.

My research group website can be found at

I am also a member of the Behaviour research group.


2006 PhD, Bristol
2003 BSc, Bristol (Zoology)


2017 -  Professor of Sensory and Evolutionary Ecology

2015 - Associate Professor of Sensory and Evolutionary Ecology

2012-2014 BBSRC David Philips Senior Research Fellow, Exeter

2009-2012 Fellow, Churchill College, Cambridge

2009-2012 BBSRC David Phillips Research Fellow, Cambridge

2006-2009 Research Fellow, Girton College, Cambridge

2006 PhD Bristol


Research group links

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Research interests

More up to date information is available on my lab website.

We study a range of subjects in sensory and evolutionary ecology, but most notably adaptive coloration and vision. Most of our research covers anti-predator coloration, behaviour, drivers of diversity, human impacts, and methods to study vision and visual signals. Our work is interdisciplinary, spanning sensory, behavioural, evolutionary biology, and incorporates ideas and methods from experimental psychology and computer vision. We work on various taxonomic groups, including birds, reptiles, crustaceans, insects, and primates. The focus of much of our current work is coloration and behaviour in marine invertebrates, including anthropogenic impacts on this such as pollution. 

Currently, two major areas of research in the lab are to study animal colour change for camouflage (both mechanisms and functions), and applied benefits of understanding animal vision.

Some of our main aims are to:

  • Understand animal coloration in terms of how the signals are perceived by the relevant receiver(s), and use this to study how visual information guides behaviour. 
  • Determine how human impacts such as pollution and invasive species are affecting animal coloration, behaviour and ecology, and how knowledge of animal vision can lead to applied benefits and help solve some of these problems.
  • Determine how arms races and coevolution can drive both intra- and interspecific diversity in animal coloration and vision.
  • Understand how the form and diversity of coloration and visual systems results from varied selection pressures, for example from different habitat structures and for both communication and predator avoidance.

Anti-Predator Coloration
Much of our research aims to understand various forms of anti-predator coloration. A major avenue of research is work on camouflage, whereby we aim to understand how different types of camouflage work (e.g. background matching, disruptive coloration), and the survival value that they bring. Much of our current work focusses on marine invertebrates such as chameleon prawns, sea slugs, anemones, and crabs. This includes studies of colour change, background choice, warning signals, and mimicry. 

A major area is to test the mechanisms and adaptive value of colour change in animals. To do so, we study a range of marine species, including crabs, chameleon prawns, and fish. This work also includes testing the influence of climate change and pollution on colour change and camouflage.

We have also had further key projects investigating camouflage in wild animals, such as ground nesting birds, especially in several species of plover in South Africa and nightjar in Zambia. 

Vision & Image Analysis Methods to Study Visual Signals
Our work on adaptive coloration is underpinned by trying to understand the visual mechanisms involved. The model receivers we are most interested in are birds, and we use and produce models analysing visual signals from a bird’s visual perspective. In addition, we have also modelled insect and primate vision, and collaborate with James Higham (New York University) on primate visual signals. We are also interested in developing techniques, especially digital image analysis, to quantify complex two dimensional signals, both in terms of pattern and colour with respect to animal vision and producing models of spatial vision.

A more recent major avaneue of work is to explore how undertsanding animal vision can have applied benefits in developing new products and informing about issues related to animal welfare and productivity.

Research Grants

In addition to those below, I have received various grants from industry.

  • 01/10/2017 - 30/09/2020: BBSRC Industrial Partnership Award (part funded by QinetiQ. How to optimise imperfect camouflage.
  • 01/09/2017 – 01/02/2018: Racing Foundation and British Horseracing Authority. Horse vision, obstacle visibility, and safety.
  • 14/11/2016 – 14/05/2017: BBSRC Pathfinder. Imaging Animal Vision.
  • 10/08/2016 – 24/08/2016: Fundação de Amparo à Pesquisa do Estado de São Paulo – FAPESP. Visiting research collaboration at University of São Paulo.
  • 01/10/2014 - 30/09/2017: BBSRC Standard Grant. Predator learning of camouflage types.
  • 01/08/2012 – 30/09/2014: BBSRC Standard Grant. Predator Vision and Avian Egg Camouflage.
  • 01/10/2009 - 30/09/2014: BBSRC David Phillips Fellowship. Predator vision and defensive coloration: from mechanism to function.
  • 2008: Royal Society Research Grant. Predator perception and the evolutionary design of warning signals.
  • A range of smaller grants and various undergraduate summer projects grants from organisations like the Nuffield Foundation, ASAB, BES, and the BBSRC.

Research networks

External Collaborators

  • Graeme Ruxton (St Andrews): Anti-predator behaviour, prey movement, and coloration.
  • James Higham (New York University): Primate vision and sexual signals.
  • Johanna Mappes (University of Jyväskylä): Predator foraging, the evolution of prey polymorphisms, and warning signals.
  • Carita Lindstedt: (University of Jyväskylä): Prey polymorphisms, and warning signals.
  • Claire Spottiswoode (Cambridge, Zoology): Coevolution and arms races in African parasitic finches and their hosts, and the evolution of egg colour polymorphism and rejection behaviour.
  • Chris Jiggins (Cambridge, Zoology): Evolution of mimicry and selection on Heliconius butterflies.
  • Keita Tanaka (Rikkyo University): Brood parasitism in cuckoos and begging signals.
  • David Tolhurst (Cambridge, Department of Physiology, Development & Neuroscience): Psychophysics and biology of anti-predator coloration and motion perception.
  • Peter Todd (National University of Singapore): Crab camouflage and polymorphism.
  • John Skelhorn (Newcastle University): Camouflage and learning.


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Stevens M (2021). Life in Colour How Animals See the World., Random House. Abstract.
Stevens M (2016). Cheats and Deceits How Animals and Plants Exploit and Mislead., Oxford University Press. Abstract.
Stevens M (2013). Sensory Ecology, Behaviour, and Evolution., OUP Oxford.
Stevens M, Merilaita S (2011). Animal Camouflage: Mechanisms and Function., Cambridge University Press.

Journal articles

Twort L, Stevens M (In Press). Active background choice facilitates camouflage in shore crabs (Carcinus maenas). Animal Behaviour
Tan M, Zhang S, Stevens M, Li D, Tan EJ (In Press). Antipredator defences in motion: animals reduce predation risks by concealing or misleading motion signals. Biological Reviews
Price N, Green S, Troscianko J, Tregenza T, Stevens M (In Press). Background matching and disruptive coloration as habitat-specific strategies for camouflage. Scientific Reports
Green S, Duarte RC, Kellett E, Alagaratnam N, Stevens M (In Press). Color change and behavioral choice facilitate chameleon prawn camouflage against different seaweed backgrounds. Communications Biology
Galloway J, Green SD, Stevens M, Kelley L (In Press). Finding a signal hidden among noise: how can predators overcome camouflage strategies?. Philosophical Transactions of the Royal Society B: Biological Sciences
Paul S, Stevens M (In Press). Horse vision and obstacle visibility in horseracing. Applied Animal Behaviour Science
Carter EE, Tregenza T, Stevens M (In Press). Ship noise inhibits colour change, camouflage, and anti-predator behaviour in shore crabs. Current Biology
Somerville J, Blount J, Stevens M (In Press). The use of vision modelling to design bycatch reduction devices using light. Fish and Fisheries
Hughes A, Briolat E, Arenas L, Liggins E, Stevens M (In Press). Varying benefits of generalist and specialist camouflage in two versus four background environments. Behavioral Ecology
Lund J, Dixit T, Attwood M, Hamama S, Moya C, Jamie G, Spottiswoode C, Stevens M (In Press). When perfection isn’t enough: host egg signatures are an effective defence against high-fidelity African cuckoo mimicry. Proceedings of the Royal Society B: Biological Sciences
Rising K, Hardege J, Tregenza T, Stevens M (2022). Anthropogenic noise may impair the mating behaviour of the Shore Crab Carcinus Maenas. PLoS One, 17(10). Abstract.  Author URL.
Niu Y, Stevens M, Sun H (2021). Commercial harvesting has driven the evolution of camouflage in an alpine plant. Current Biology, 31, 446-449.
Duarte RC, Dias GM, Flores AAV, Stevens M (2021). Different ontogenetic trajectories of body colour, pattern and crypsis in two sympatric intertidal crab species. Biological Journal of the Linnean Society, 132(1), 17-31. Abstract.
Briolat ES, Arenas LM, Hughes AE, Liggins E, Stevens M (2021). Generalist camouflage can be more successful than microhabitat specialisation in natural environments. BMC Ecol Evol, 21(1). Abstract.  Author URL.
Caves EM, Dixit T, Colebrook-Robjent JFR, Hamusikili L, Stevens M, Thorogood R, Spottiswoode CN (2021). Hosts elevate either within-clutch consistency or between-clutch distinctiveness of egg phenotypes in defence against brood parasites. Proc Biol Sci, 288(1953). Abstract.  Author URL.
Troscianko J, Nokelainen O, Skelhorn J, Stevens M (2021). Variable crab camouflage patterns defeat search image formation. Commun Biol, 4(1). Abstract.  Author URL.
Spaniol RL, Mendonça MDS, Hartz SM, Iserhard CA, Stevens M (2020). Discolouring the Amazon Rainforest: how deforestation is affecting butterfly coloration. Biodiversity and Conservation, 29(9-10), 2821-2838.
Stephenson JF, Stevens M, Troscianko J, Jokela J (2020). The Size, Symmetry, and Color Saturation of a Male Guppy's Ornaments Forecast His Resistance to Parasites. AMERICAN NATURALIST, 196(5), 597-608.  Author URL.
Green SD, Duarte RC, Kellett E, Alagaratnam N, Stevens M (2019). Colour change and behavioural choice facilitate chameleon prawn camouflage against different seaweed backgrounds. Commun Biol, 2(1). Abstract.  Author URL.
Narasimha S, Nagornov KO, Menin L, Mucciolo A, Rohwedder A, Humbel BM, Stevens M, Thum AS, Tsybin YO, Vijendravarma RK, et al (2019). Drosophila melanogaster cloak their eggs with pheromones, which prevents cannibalism. PLoS Biology, 17
Stoddard MC, Hogan BG, Stevens M, Spottiswoode CN (2019). Higher-level pattern features provide additional information to birds when recognizing and rejecting parasitic eggs. Philosophical Transactions of the Royal Society B: Biological Sciences, 374(1769). Abstract.
Hughes A, Liggins E, Stevens M (2019). Imperfect camouflage: how to hide in a variable world?. Proceedings of the Royal Society B: Biological Sciences, 286, 20190646-20190646.
Nokelainen O, Maynes R, Mynott S, Price N, Stevens M (2019). Improved camouflage through ontogenetic colour change confers reduced detection risk in shore crabs. Functional Ecology, 33(4), 654-669. Abstract.
Briolat E, Zagrobelny M, Olsen C, Blount J, Stevens M (2019). No evidence of quantitative signal honesty across species of aposematic burnet moths (Lepidoptera: Zygaenidae). Journal of Evolutionary Biology, 32, 31-48.
Nokelainen O, Maynes R, Mynott S, Price N, Stevens M (2019). Supplementary data for: Improved camouflage through ontogenetic colour change confers reduced detection risk in shore crabs. JYX
Dell’Aglio DD, Troscianko J, Stevens M, McMillan WO, Jiggins CD (2019). The conspicuousness of the toxic<i>Heliconius</i>butterflies across time and habitat. Abstract.
Stevens M, Ruxton GD (2019). The key role of behaviour in animal camouflage. Biological Reviews, 94(1), 116-134. Abstract.
Chan IZW, Stevens M, Todd PA (2019). pat-geom: a software package for the analysis of animal patterns. Methods in Ecology and Evolution, 10(4), 591-600. Abstract.
Walton OC, Stevens M (2018). Avian vision models and field experiments determine the survival value of peppered moth camouflage. Communications Biology, 1, 118-118.
Troscianko J, Skelhorn J, Stevens M (2018). Camouflage strategies interfere differently with observer search images. Proceedings of the Royal Society B: Biological Sciences, 285(1886). Abstract.
Troscianko J, Skelhorn J, Stevens M (2018). Camouflage strategies interfere differently with observer search images. Proceedings of the Royal Society B: Biological Sciences, 285, 20181386-20181386.
Badás EP, Martínez J, Rivero-de Aguilar J, Ponce C, Stevens M, Merino S (2018). Colour change in a structural ornament is related to individual quality, parasites and mating patterns in the blue tit. Science of Nature, 105(1-2). Abstract.
Nokelainen O, Stevens M, Caro T (2018). Colour polymorphism in the coconut crab (Birgus latro). Evolutionary Ecology, 32, 75-75.
Ligon RA, Diaz CD, Morano JL, Troscianko J, Stevens M, Moskeland A, Laman TG, Scholes E (2018). Evolution of correlated complexity in the radically different courtship signals of birds-of-paradise. PLoS Biology, 16, e2006962-e2006962.
Gómez J, Ramo C, Troscianko J, Stevens M, Castro M, Pérez-Hurtado A, Liñán-Cembrano G, Amat JA (2018). Individual egg camouflage is influenced by microhabitat selection and use of nest materials in ground-nesting birds. Behavioral Ecology and Sociobiology, 72, 142-142.
Paul SC, Stevens M, Burton J, Pell JK, Birkett MA, Blount JD (2018). Invasive egg predators and food availability interactively affect maternal investment in egg chemical defense. Frontiers in Ecology and Evolution, 6(JAN). Abstract.
Gómez J, Ramo C, Stevens M, Liñán-Cembrano G, Rendón MA, Troscianko J, Amat JA (2018). Latitudinal variation in biophysical characteristics of avian eggshells to cope with differential effects of solar radiation. Ecology and Evolution, 8, 8019-8029.
Paul SC, Stevens M, Pell JK, Birkett MA, Blount JD (2018). Parental phenotype not predator cues influence egg warning coloration and defence levels. Animal Behaviour, 140, 177-186. Abstract.
Niu Y, Sun H, Stevens M (2018). Plant camouflage: ecology, evolution, and implications. Trends in Ecology and Evolution, 33, 608-618.
Smithers SP, Rooney R, Wilson A, Stevens M (2018). Rock pool fish use a combination of colour change and substrate choice to improve camouflage. Animal Behaviour, 144, 53-65. Abstract.
Briolat E, Zagrobelny M, Olsen, CE, Blount J, Stevens M (2018). Sex differences but no evidence of quantitative honesty in the warning signals of six-spot burnet moths (Zygaena filipendulae L.). Evolution, 77, 1460-1474.
Duarte RC, Stevens M, Flores AAV (2018). The adaptive value of camouflage and colour change in a polymorphic prawn. Scientific Reports, 8, 16028-16028.
Dell'Aglio DD, Troscianko J, McMillan WO, Stevens M, Jiggins CD (2018). The appearance of mimetic Heliconius butterflies to predators and conspecifics. Evolution, 72(10), 2156-2166. Abstract.
Mynott S, Daniels C, Widdicombe S, Stevens M (2018). Using camouflage for conservation: colour change in juvenile European lobster. Abstract.
Duarte RC, Flores AAV, Stevens M (2017). Camouflage through colour change: mechanisms, adaptive value, and ecological significance. Philosophical Transactions B: Biological Sciences, 372, 20160342-20160342.
Niu Y, Chen Z, Stevens M, Sun H (2017). Divergence in cryptic leaf colour provides local camouflage in an alpine plant. Proceedings of the Royal Society B: Biological Sciences, 284, 20171654-20171654.
Arenas LM, Stevens M (2017). Diversity in warning coloration is easily recognised by avian predators. Journal of Evolutionary Biology, 30, 1288-1302.
Caves EM, Stevens M, Spottiswoode CN (2017). Does coevolution with a shared parasite drive hosts to partition their defences among species?. Proceedings of the Royal Society B: Biological Sciences, 284(1854), 20170272-20170272. Abstract.
Geltsch N, Moskat C, Elek Z, Ban M, Stevens M (2017). Egg spotting pattern in the common cuckoo and its great reed warbler host: a century perspective. Biological Journal of the Linnean Society, 121, 50-50.
Badás EP, Martínez J, Rivero-de Aguilar J, Stevens M, van der Velde M, Komdeur J, Merino S (2017). Eggshell pigmentation in the blue tit: male quality matters. Behavioral Ecology and Sociobiology, 71, 57-57.
Stevens M, Troscianko J, Wilson-Aggarwal JK, Spottiswoode CN (2017). Improvement of individual camouflage through background choice in ground-nesting birds. Nature Ecology and Evolution, 1, 1325-1325.
Koski T-M, Lindstedt C, Klemola T, Troscianko J, Mäntylä E, Tyystjärvi E, Stevens M, Helander M, Laaksonen T (2017). Insect herbivory may cause changes in the visual properties of leaves and affect the camouflage of herbivores to avian predators. Behavioral Ecology and Sociobiology, 31, 97-97.
Troscianko J, Skelhorn J, Stevens M (2017). Quantifying camouflage: how to predict detectability from appearance. BMC Evolutionary Biology, 17, 7-7.
Troscianko J, Wilson-Aggarwal J, Griffiths D, Spottiswoode CN, Stevens M (2017). Relative advantages of dichromatic and trichromatic color vision in camouflage breaking. Behavioral Ecology, 28, 556-556.
Smithers SP, Wilson A, Stevens M (2017). Rock pool gobies change their body pattern in response to background features. Biological Journal of the Linnean Society, 121, 109-121.
Pérez-Rodríguez L, Jovani R, Stevens M (2017). Shape matters: animal colour patterns as signals of individual quality. Proceedings of the Royal Society B: Biological Sciences, 284, 20162446-20162446.
Cuthill IC, Allen WL, Arbuckle K, Caspers B, Chaplin G, Hauber ME, Hill GE, Jablonski NG, Jiggins CD, Kelber A, et al (2017). The biology of color. Science, 357(6350), eaan0221-eaan0221. Abstract.
Nokelainen O, Hubbard N, Lown AE, Wood LE, Stevens M (2017). Through predators’ eyes - phenotype-environment associations in shore crab coloration at different spatial scales. Biological Journal of the Linnean Society, 122, 738-751.
Dell'aglio DD, Stevens M, Jiggins CD (2016). Avoidance of an aposematically coloured butterfly by wild birds in a tropical forest. Ecological Entomology, 41(5), 627-632. Abstract.
Medina I, Troscianko J, Stevens M, Langmore NE (2016). Brood Parasitism is Linked to Egg Pattern Diversity within and among Species of Australian Passerines. Am Nat, 187(3), 351-362. Abstract.  Author URL.
Troscianko J, Wilson-Aggarwal J, Stevens M, Spottiswoode CN (2016). Camou age predicts survival in ground-nesting birds. Scientific Reports, 6
Nokelainen O, Stevens M (2016). Camouflage. Current Biology, 26(14), R654-R656. Abstract.
Stevens M (2016). Color Change, Phenotypic Plasticity, and Camouflage. Frontiers in Ecology and Evolution, 4
Wilson-Aggarwal, Troscianko, Stevens M, Spottiswoode CN (2016). Escape distance in ground-nesting birds differs with individual level of camouflage. The American Naturalist, 188, 231-231.
Marshall KLA, Philpott KE, Stevens M (2016). Microhabitat choice in island lizards enhances camou age against avian predators. Scientific Reports, 6
Troscianko, Wilson-Aggarwal, Spottiswoode CN, Stevens M (2016). Nest covering in plovers: how modifying the visual environment influences egg camouflage. Ecology and Evolution, 6, 7536-7545.
Chan IZW, Stevens M, Todd PA (2016). Quantifying shell pattern and colour polymorphism in the button snail Umbonium vestiarium (Mollusca: Gastropoda: Trochacea) and comparing morph frequencies between two populations using the Mantel test. Raffles Bulletin of Zoology, 2016, 22-32. Abstract.
Duarte RC, Stevens M, Flores AAV (2016). Shape, colour plasticity, and habitat use indicate morph-specific camouflage strategies in a marine shrimp. BMC Evolutionary Biology, 16, 218-218.
Caro T, Sherratt TN, Stevens M (2016). The ecology of multiple colour defences. Evolutionary Ecology, 30, 797-809.
Stevens M (2016). When to attack defended prey? a comment on Skelhorn et al. Behavioral Ecology, 27(4).
Stevens M (2015). Anti-Predator Coloration and Behaviour: a Longstanding Topic with Many Outstanding Questions. CURRENT ZOOLOGY, 61(4), 702-707.  Author URL.
Flores EE, Stevens M, Moore AJ, Rowland HM, Blount JD (2015). Body size but not warning signal luminance influences predation risk in recently metamorphosed poison frogs. Ecology and evolution, 5(20), 4603-4616. Abstract.
Burriss R, Troscianko J, Lovell PG, Fulford AJC, Stevens M, Quigley R, Payne J, Saxton TK, Rowland HM (2015). Changes in Women’s Facial Skin Color over the Ovulatory Cycle are Not Detectable by the Human Visual System. PLoS ONE, 10, e0130093-e0130093.
Marshall KLA, Philpot KE, Stevens M (2015). Conspicuous male coloration impairs survival against avian predators in Aegean wall lizards, Podarcis erhardii. Ecology and evolution, 5(18), 4115-4131. Abstract.
Troscianko J, Stevens M, Skelhorn J (2015). Does Disruptive Camouflage Disrupt Search Image Formation?. PERCEPTION, 44(10), 1239-1240.  Author URL.
Stevens M (2015). Evolutionary Ecology: Insect Mothers Control Their Egg Colours. Curr Biol, 25(17), R755-R757. Abstract.  Author URL.
Caves EM, Stevens M, Iversen ES, Spottiswoode CN (2015). Hosts of avian brood parasites have evolved egg signatures with elevated information content. Proc Biol Sci, 282(1810). Abstract.  Author URL.
Troscianko J, Stevens M (2015). Image Calibration and Analysis Toolbox – a free software suite for objectively measuring reflectance, colour and pattern. Methods in Ecology and Evolution, 6, 1320-1331.
Marshall KLA, Philpot KE, Damas-Moreira I, Stevens M (2015). Intraspecific Colour Variation among Lizards in Distinct Island Environments Enhances Local Camouflage. PLoS One, 10(9). Abstract.  Author URL.
Stevens M, Broderick AC, Godley BJ, Lown AE, Troscianko J, Weber N, Weber SB (2015). Phenotype-environment matching in sand fleas. Biol Lett, 11(8). Abstract.  Author URL.
Arenas LM, Walter D, Stevens M (2015). Signal honesty and predation risk among a closely related group of aposematic species. Scientific Reports, 5, 11021-11021.
Ruxton GD, Stevens M (2015). The evolutionary ecology of decorating behaviour. Biol Lett, 11(6). Abstract.  Author URL.
Hughes AE, Magor-Elliott RS, Stevens M (2015). The role of stripe orientation in target capture success. Frontiers in Zoology, 12, 17-17.
Stevens M (2015). Visual Ecology. Marine and Freshwater Behaviour and Physiology, 48(3), 221-223.
Stevens M, Lown AE, Wood LE (2014). Camouflage and individual variation in shore crabs (Carcinus maenas) from different habitats. PLoS One, 9(12). Abstract.  Author URL.
Kang C, Stevens M, Moon J-Y, Lee S-I, Jablonski PG (2014). Camouflage through behavior in moths: the role of background matching and disruptive coloration. Behavioral Ecology
Allen WL, Stevens M, Higham JP (2014). Character displacement of Cercopithecini primate visual signals. Nat Commun, 5 Abstract.  Author URL.
Arenas LM, Troscianko J, Stevens M (2014). Color contrast and stability as key elements for effective warning signals. Frontiers in Ecology and Evolution, 2
Stevens M, Lown AE, Wood LE (2014). Colour change and camouflage in juvenile shore crabs Carcinus maenas. Frontiers in Ecology and Evolution, 2
Stevens M (2014). Confusion and illusion: understanding visual traits and behavior. A comment on Kelley and Kelley. Behavioral Ecology, 25(3), 464-465.
Stevens M, Ruxton GD (2014). Do animal eyespots really mimic eyes?. Current Zoology, 60(1), 26-36. Abstract.
Stevens M (2014). Evolution: predator versus parasite. Curr Biol, 24(10), R388-R390. Abstract.  Author URL.
Winters AE, Stevens M, Mitchell C, Blomberg SP, Blount JD (2014). Maternal effects and warning signal honesty in eggs and offspring of an aposematic ladybird beetle. Functional Ecology, 28(5), 1187-1196. Abstract.
Hughes AE, Stevens M, Tolhurst DJ (2014). Methods and mechanisms of motion dazzle. I-PERCEPTION, 5(5).  Author URL.
Hughes AE, Troscianko J, Stevens M (2014). Motion dazzle and the effects of target patterning on capture success. BMC Evolutionary Biology, 14
Hughes AE, Troscianko J, Stevens M (2014). Motion dazzle and the effects of target patterning on capture success. BMC Evol Biol, 14 Abstract.  Author URL.
Stevens M, Lown AE, Denton AM (2014). Rockpool gobies change colour for camouflage. PLoS One, 9
Marshall KLA, Stevens M (2014). Wall lizards display conspicuous signals to conspecifics and reduce detection by avian predators. Behavioral Ecology
(2013). A window on the past: male ornamental plumage reveals the quality of their early-life environment. Proceedings. Biological sciences / the Royal Society, 280(1756). Abstract.
Walker LK, Stevens M, Karadaş F, Kilner RM, Ewen JG (2013). A window on the past: male ornamental plumage reveals the quality of their early-life environment. Proc Biol Sci, 280(1756). Abstract.  Author URL.
Stevens M (2013). Bird brood parasitism. Curr Biol, 23(20), R909-R913. Abstract.  Author URL.
Stevens M, Rong CP, Todd PA (2013). Colour change and camouflage in the horned ghost crab Ocypode ceratophthalmus. Biological Journal of the Linnean Society, 109(2), 257-270. Abstract.
Stevens M (2013). Concealing Coloration in Animals. Animal Behaviour, 86(6), 1333-1334.
Troscianko J, Lown AE, Hughes AE, Stevens M (2013). Defeating crypsis: detection and learning of camouflage strategies. PLoS One, 8(9). Abstract.  Author URL.
Flores EE, Stevens M, Moore AJ, Blount JD (2013). Diet, development and the optimization of warning signals in post-metamorphic green and black poison frogs. Functional Ecology
Flores EE, Stevens M, Moore AJ, Blount JD (2013). Diet, development and the optimization of warning signals in post-metamorphic green and black poison frogs. Functional Ecology, 27(3), 816-829. Abstract.
Teasdale LC, Stevens M, Stuart-Fox D (2013). Discrete colour polymorphism in the tawny dragon lizard (Ctenophorus decresii) and differences in signal conspicuousness among morphs. Journal of Evolutionary Biology
Teasdale LC, Stevens M, Stuart-Fox D (2013). Discrete colour polymorphism in the tawny dragon lizard (Ctenophorus decresii) and differences in signal conspicuousness among morphs. Journal of Evolutionary Biology, 26(5), 1035-1046. Abstract.
Stevens M (2013). Evolutionary ecology: knowing how to hide your eggs. Curr Biol, 23(3), R106-R108. Abstract.  Author URL.
Stevens M (2013). Exchanging messages between plants and animals. Trends in Ecology & Evolution, 28(7), 386-387.
Stevens M, Troscianko J, Spottiswoode CN (2013). Repeated targeting of the same hosts by a brood parasite compromises host egg rejection. Nat Commun, 4 Abstract.  Author URL.
Stevens M, Marshall KLA, Troscianko J, Finlay S, Burnand D, Chadwick SL (2013). Revealed by conspicuousness: Distractive markings reduce camouflage. Behavioral Ecology, 24(1), 213-222. Abstract.
Higham JP, Pfefferle D, Heistermann M, Maestripieri D, Stevens M (2013). Signaling in multiple modalities in male rhesus macaques: sex skin coloration and barks in relation to androgen levels, social status, and mating behavior. BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY, 67(9), 1457-1469.  Author URL.
Higham JP, Pfefferle D, Heistermann M, Maestripieri D, Stevens M (2013). Signaling in multiple modalities in male rhesus macaques: sex skin coloration and barks in relation to androgen levels, social status, and mating behavior. Behavioral Ecology and Sociobiology, 1-13.
Stevens M, Troscianko J, Marshall KLA, Finlay S (2013). What is camouflage through distractive markings a reply to Merilaita et al. (2013). Behavioral Ecology, 24(5).
Merrill RM, Wallbank RWR, Bull V, Salazar PCA, Mallet J, Stevens M, Jiggins CD (2012). Disruptive ecological selection on a mating cue. Proc Biol Sci, 279(1749), 4907-4913. Abstract.  Author URL.
Hughes A, Stevens M, Tolhurst D (2012). Factors that affect tracking of occluded target motion. PERCEPTION, 41, 118-118.  Author URL.
Spottiswoode CN, Stevens M (2012). Host-Parasite Arms Races and Rapid Changes in Bird Egg Appearance. AMERICAN NATURALIST, 179(5), 633-648.  Author URL.
Spottiswoode CN, Stevens M (2012). Host-parasite arms races and rapid changes in bird egg appearance. Am Nat, 179(5), 633-648. Abstract.  Author URL.
Stevens M, Ruxton GD (2012). Linking the evolution and form of warning coloration in nature. Proc Biol Sci, 279(1728), 417-426. Abstract.  Author URL.
Hughes A, Troscianko J, Tolhurst D, Stevens M (2012). Prey movement and the mechanisms of dazzle coloration. PERCEPTION, 41(10), 1271-1271.  Author URL.
Stevens M (2011). Avian vision and egg colouration: Concepts and measurements. Avian Biology Research, 4(4), 168-184. Abstract.
Stoddard MC, Stevens M (2011). Avian vision and the evolution of egg color mimicry in the common cuckoo. Evolution, 65(7), 2004-2013. Abstract.  Author URL.
Lindstedt C, Eager H, Ihalainen E, Kahilainen A, Stevens M, Mappes J (2011). Direction and strength of selection by predators for the color of the aposematic wood tiger moth. Behavioral Ecology, 22(3), 580-587. Abstract.
Higham JP, Hughes KD, Brent LJN, Dubuc C, Engelhardt A, Heistermann M, Maestriperi D, Santos LR, Stevens M (2011). Familiarity affects the assessment of female facial signals of fertility by free-ranging male rhesus macaques. Proceedings of the Royal Society B: Biological Sciences, 278(1723), 3452-3458. Abstract.
Spottiswoode CN, Stevens M (2011). How to evade a coevolving brood parasite: egg discrimination versus egg variability as host defences. Proc Biol Sci, 278(1724), 3566-3573. Abstract.  Author URL.
Stevens M, Searle WTL, Seymour JE, Marshall KLA, Ruxton GD (2011). Motion dazzle and camouflage as distinct anti-predator defenses. BMC Biol, 9 Abstract.  Author URL.
Tanaka KD, Morimoto G, Stevens M, Ueda K (2011). Rethinking visual supernormal stimuli in cuckoos: Visual modeling of host and parasite signals. Behavioral Ecology, 22(5), 1012-1019. Abstract.
Langmore NE, Stevens M, Maurer G, Heinsohn R, Hall ML, Peters A, Kilner RM (2011). Visual mimicry of host nestlings by cuckoos. Proceedings of the Royal Society B: Biological Sciences, 278(1717), 2455-2463. Abstract.
Higham JP, Brent LJN, Dubuc C, Accamando AK, Engelhardt A, Gerald MS, Heistermann M, Stevens M (2010). Color signal information content and the eye of the beholder: a case study in the rhesus macaque. Behavioral Ecology, 21(4), 739-746. Abstract.
Stoddard MC, Stevens M (2010). Pattern mimicry of host eggs by the common cuckoo, as seen through a bird's eye. Proc Biol Sci, 277(1686), 1387-1393. Abstract.  Author URL.
Stevens M (2010). Sensory ecology, evolution, and behavior. Current Zoology, 56(3).
Shultziner D, Stevens T, Stevens M, Stewart BA, Hannagan RJ, Saltini-Semerari G (2010). The causes and scope of political egalitarianism during the Last Glacial: a multi-disciplinary perspective. Biology and Philosophy, 25(3), 319-346. Abstract.
Sandre SL, Stevens M, Mappes J (2010). The effect of predator appetite, prey warning coloration and luminance on predator foraging decisions. Behaviour, 147(9), 1121-1143. Abstract.
Spottiswoode CN, Stevens M (2010). Visual modeling shows that avian host parents use multiple visual cues in rejecting parasitic eggs. Proc Natl Acad Sci U S A, 107(19), 8672-8676. Abstract.  Author URL.
Stevens M, Merilaita S (2009). Animal camouflage: current issues and new perspectives. Philos Trans R Soc Lond B Biol Sci, 364(1516), 423-427.  Author URL.
Langmore NE, Stevens M, Maurer G, Kilner RM (2009). Are dark cuckoo eggs cryptic in host nests?. Animal Behaviour, 78(2), 461-468. Abstract.
Stevens M, Merilaita S (2009). Defining disruptive coloration and distinguishing its functions. Philos Trans R Soc Lond B Biol Sci, 364(1516), 481-488. Abstract.  Author URL.
Stevens M, Winney IS, Cantor A, Graham J (2009). Outline and surface disruption in animal camouflage. Proc Biol Sci, 276(1657), 781-786. Abstract.  Author URL.
Stevens M, Stoddard MC, Higham JP (2009). Studying primate color: Towards visual system-dependent methods. International Journal of Primatology, 30(6), 893-917. Abstract.
Stevens M, Cantor A, Graham J, Winney IS (2009). The function of animal 'eyespots': Conspicuousness but not eye mimicry is key. Current Zoology, 55(5), 319-326. Abstract.
Stevens M, Castor-Perry SA, Price JRF (2009). The protective value of conspicuous signals is not impaired by shape, size, or position asymmetry. Behavioral Ecology, 20(1), 96-102. Abstract.
Stevens M, Hardman CJ, Stubbins CL (2008). Conspicuousness, not eye mimicry, makes "eyespots" effective antipredator signals. Behavioral Ecology, 19(3), 525-531. Abstract.
Stevens M, Yule DH, Ruxton GD (2008). Dazzle coloration and prey movement. Proc Biol Sci, 275(1651), 2639-2643. Abstract.  Author URL.
Stevens M, Graham J, Winney IS, Cantor A (2008). Testing Thayer's hypothesis: can camouflage work by distraction?. Biol Lett, 4(6), 648-650. Abstract.  Author URL.
Stevens M, Stubbins CL, Hardman CJ (2008). The anti-predator function of 'eyespots' on camouflaged and conspicuous prey. Behavioral Ecology and Sociobiology, 62(11), 1787-1793. Abstract.
Houston AI, Stevens M, Cuthill IC (2007). Animal camouflage: Compromise or specialize in a 2 patch-type environment?. Behavioral Ecology, 18(4), 769-775. Abstract.
Rowland HM, Speed MP, Ruxton GD, Edmunds M, Stevens M, Harvey IF (2007). Countershading enhances cryptic protection: an experiment with wild birds and artificial prey. Animal Behaviour, 74(5), 1249-1258. Abstract.
Cuthill IC, Troscianko TS, Kibblewhite A, King O, Stevens M (2007). Edge enhancement in disruptive camouflage. PERCEPTION, 36(9), 1399-1399.  Author URL.
Stevens M, Hopkins E, Hinde W, Adcock A, Connolly Y, Troscianko T, Cuthill IC (2007). Field experiments on the effectiveness of 'eyespots' as predator deterrents. Animal Behaviour, 74(5), 1215-1227. Abstract.
Stevens M, Cuthill IC (2007). Hidden messages: Are ultraviolet signals a special channel in avian communication?. BioScience, 57(6), 501-507. Abstract.
Stevens M (2007). Predator perception and the interrelation between different forms of protective coloration. Proc Biol Sci, 274(1617), 1457-1464. Abstract.  Author URL.
Stevens M, Párraga CA, Cuthill IC, Partridge JC, Troscianko TS (2007). Using digital photography to study animal coloration. Biological Journal of the Linnean Society, 90(2), 211-237. Abstract.
Stevens M, Cuthill IC (2006). Disruptive coloration, crypsis and edge detection in early visual processing. Proc Biol Sci, 273(1598), 2141-2147. Abstract.  Author URL.
Stevens M, Cuthill IC, Windsor AMM, Walker HJ (2006). Disruptive contrast in animal camouflage. Proc Biol Sci, 273(1600), 2433-2438. Abstract.  Author URL.
Stevens M, Cuthill IC, Alejandro Párraga C, Troscianko T (2006). The effectiveness of disruptive coloration as a concealment strategy. Prog Brain Res, 155, 49-64. Abstract.  Author URL.
Cuthill IC, Stevens M, Windsor AMM, Walker HJ (2006). The effects of pattern symmetry on detection of disruptive and background matching coloration. Behavioral Ecology, 17(5), 828-832.
Troscianko T, Cuthill IC, Stevens M, Graham L, Richardson S, Párraga CA (2005). Detection of cryptic targets in avian vision:: a field study. PERCEPTION, 34, 47-47.  Author URL.
Cuthill IC, Stevens M, Sheppard J, Maddocks T, Párraga CA, Troscianko TS (2005). Disruptive coloration and background pattern matching. Nature, 434(7029), 72-74. Abstract.  Author URL.
Stevens M (2005). The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera. Biol Rev Camb Philos Soc, 80(4), 573-588. Abstract.  Author URL.
Stevens M, Cuthill IC (2005). The unsuitability of HTML-based colour charts for estimating animal colours--a comment on Berggren and Merilä (2004). Front Zool, 2 Abstract.  Author URL.
Stevens M, Cuthill IC, Sheppard J, Maddocks T, Troscianko T, Párraga CA (2004). Disruptive colouration and background pattern matching in insect crypsis. PERCEPTION, 33, 35-36.  Author URL.
Franks NR, Dornhaus A, Fitzsimmons JP, Stevens M (2003). Speed versus accuracy in collective decision making. Proc Biol Sci, 270(1532), 2457-2463. Abstract.  Author URL.


Stevens M, Arenas LM, Lown AE (2016). Color in camouflage, mimicry, and warning signals. In  (Ed) Handbook of Color Psychology, Cambridge University Press (CUP), 357-376.
Stevens M (2013). Sensing the World. In  (Ed) Sensory Ecology, Behaviour, and Evolution, Oxford University Press, 21-39.
Stevens M (2013). Sensory Ecology, Behaviour, and Evolution. In Stevens M (Ed) Sensory Ecology, Behaviour, and Evolution, OUP Oxford.
Stevens M (2013). Sensory Ecology, Information, and Decision-Making. In  (Ed) Sensory Ecology, Behaviour, and Evolution, Oxford University Press, 2-18.
Stevens M, Merilaita S (2011). Animal Camouflage: an Introduction. In Stevens M, Merilaita S (Eds.) Animal Camouflage: Mechanisms & Function, Cambridge University Press, 1-16.
Stevens M (2011). Animal Camouflage: Mechanisms & Function. In Stevens M, Merilaita S (Eds.) Animal Camouflage: Mechanisms & Function, Cambridge University Press. Abstract.
Merilaita S, Stevens M (2011). Crypsis Through Background Matching. In Stevens M, Merilaita S (Eds.) Animal Camouflage, Cambridge University Press, 17-33.
Mappes J, Stevens M (2010). Information Use and Sensory Ecology. In Westneat D, Fox C (Eds.) Evolutionary Behavioral Ecology, OUP USA, 148-161. Abstract.
Caro T, Merilaita S, Stevens M (2008). The Colours of Animals: from Wallace to the Present Day. I. Cryptic Colouration. In Smith CH, Beccaloni G (Eds.) Natural Selection and Beyond: the Intellectual Legacy of Alfred Russel Wallace, OUP Oxford, 125-143. Abstract.

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External Engagement and Impact

Committee/panel activities

Director of Business, Engagement, and Innovation for CEC (2022-)

CEC Senior Management Group (2022-)

CEC Research Committtee

Exeter University, BBSRC Strategy Group (-2020)

Director of Impact, Bioscience Penryn Campus (2015-2018).

Board of Directors for the Zoological Lighting Institute (2016-2018)

BBSRC Pool of Experts (for assessing grant applications; 2014-2020)

BBSRC Schools Regional Champion for the SW (2014/15)

Nuffield Foundation Undergraduate Bursary panel member (2010 & 2011).

Memberships: Association for the Study of Animal Behaviour (ASAB); Applied Vision Association (AVA); International Primatological Society (IPS); International Society for Behavioral Ecology (ISBE).

Regular reviewer for a wide range of science, biology, sensory, and behaviour journals (more than 35), including Science, Current Biology, PNAS, Evolution, Proceedings B, American Naturalist, and Journal of Experimental Biology.

I was named as a ‘Top Referee’ for 2008, 2011, and 2012 for Proceedings of the Royal Society, B.

Editorial responsibilities

I am an Editor for Current Zoology, Frontiers in Ecology and Evolution, and BMC Zoology.

Guest Editor Philosophical Transactions of the Royal Society, Series B: ‘Animal Camouflage: Current Issues and New Perspectives’ (2009, Volume: 364, Issue: 1516), involving contributions from biologists, computer scientists, psychologists, and art historians.

I have edited a book: ‘Animal Camouflage: From Mechanisms to Function’, 2011, Cambridge University Press.

Guest Editor Current Zoology: ‘Sensory Ecology, Evolution, and Behaviour’ (2010, volume 56, issue 3), involving contributions from leading sensory ecologists working on a range of subjects and sensory modalities.

Invited lectures

A range of public lectures including at the Royal Institution, Hay Festival, Royal Geographic Society, and more.

UK: University of Sussex (Biology & Environmental Science); University of Exeter (Psychology); Royal Holloway (Psychology); Newcastle University (Neuroscience); University of Oxford (Zoology, EGI); University of Edinburgh (Institute for Evolutionary Biology); University of Bristol (Vision Institute and School of Biological Sciences); University of Bath (Biology & Biochemistry); International Primatological Conference, Edinburgh (2008); European Meeting of PhD Students in Evolutionary Biology (2013), University of Exeter.

Mainland Europe: Stockholm University (Zoology) Sweden; University of Jyväskylä (Biological and Environmental Sciences), Finland; Uppsala University (Ecology & Evolution), Sweden; Bern University (Institute of Ecology & Evolution), Switzerland; European Congress on Behavioural Biology, Dijon, France (2008); European Society for Evolutionary Biology, Lisbon, Portugal (2013); University of Geneva; Wiko, Berlin (2016).

USA: Wake Forest University (Biology); University of Nebraska (Biology); University of Chicago (Ecology and Evolution); Stanford University (Biology); Princeton University (Ecology & Evolutionary Biology); University of California Davis (Biology); University of California Santa Cruz (Biology); University of California LA (Biology); New York University (Anthropology); Hunter College, City University New York (Psychology), Smithsonian/Army and Navy Club, Washington DC, USA.

Asia: National University of Singapore (Biology x 2); International Primatological Conference, Kyoto, Japan (2010); Integrative Behavioral Biology, Xi’an, China (2011, Keynote); International Symposium on Avian Brood Parasitism, Hainan, China (2012). International Ornithological Congress, Tokyo, Japan; Australasian Society for the Study of Animal Behaviour, Brisbane (plenary, 2018).

South America: Centre for Marine Biology, University of São Paulo

Media Coverage

My work has been covered in a wide range of media including on various occasions on BBC Earth News, the New York Times, the Times, LA Times, Japan Times, USA Today, the Telegraph, the Daily Mail, the Independent, the Australian, Time Magazine, New Scientist, National Geographic, MSNBC, BBC Radio 5 Live, BBC Radio Wales, BBC Radio Scotland, CBC Radio Canada, NPR Radio USA, German Radio WDR 5, Discovery, Nature, Science, TREE, Proceedings B, Discover, Natural History Magazine, Discovery Canada, Nature News, plus a wide range of national newspapers around the world and internet sites.

TV appearances: BBC 2, Inside the Animal Mind, programme 1 (28 January 2014); BBC1 The One Show (25 February 2014); National Geographic, Jurassic CSI, programme 1 (2011).

I have also been contacted by and appeared in a wide range of other reactive media outlets, including radio programmes of the American Association for the Advancement of Science, CBC Quirks and Quarks, and NPR on topics such as why zebras have stripes and animal camouflage and behaviour. I have also advised a range of organisations including the BBC1 and BBC 2, Channel 4, PBS, and the History Channel on TV programmes, and for other publications such as BBC Gardeners' World, Smithsonian and various natural history magazines. Some of my research methods have been used in interactive museum exhibitions and school teaching in the USA. I have also helped with art exhibitions related to animal coloration.

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I and module coordinator and deliver the third year Sensory Ecology course (BIO3410). I am also module coordinator for the Global Challenges MSci field course. I have signficantly contributed to the second year Biology of Birds module (BIO2439), and teach on the Cornwall Grand Challenges fieldcourse. I also have various undergraduate tutees, and both undergraduate and MSc project students.

I am a Fellow of the Higher Education Academy.

I have written a textbook: Stevens, M. Sensory Ecology, Behaviour, and Evolution. 2013. Oxford University Press. This covers key topics in sensory ecology from an evolutionary and behavioural perspective.

I have delivered lecture modules at the University of Cambridge and was Director of Studies for Churchill College Natural Science students. In addition to departmental teaching, I have given guest lectures to students in National University of Singapore, Shaanxi Normal University China, New York University, and City University New York.

Office hours: Tuesdays and Wednesdays, 9-10 am. Contact me for a MS Teams link.



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Supervision / Group

Postdoctoral researchers

  • Anna Hughes
  • Yang Niu

Postgraduate researchers

  • Emmanuelle Briolat BBSRC funded PhD student working on warning signals in moths
  • Emily Carter
  • Jim Galloway
  • Sam Green PhD student
  • Charlotte Jeffers
  • Javier Antonio Medel
  • Sara Mynott NERC funded PhD student working on climate change and camouflage in marine animals
  • Tasha Price
  • Jenna Proctor
  • Alexandra Török BBSRC funded PhD student working on the functional and mechanistic basis of startle and flash displays in animals as anti-predator defences.
  • Maria Watson

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