Prof Martin Stevens 

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Group Leader

Professor of Sensory and Evolutionary Ecology

SERSF Building (room 1.21), Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Penryn, Cornwall, TR10 9FE. UK.

More About Martin


2017 Professor of Sensory and Evolutionary Ecology

2015 Associate Professor of Sensory and Evolutionary Ecology

2012-2014 BBSRC David Philips Senior Research Fellow

2009-2012 Fellow, Churchill College, Cambridge

2009-2012 BBSRC David Phillips Senior Research Fellow, Cambridge

2006-2009 Research Fellow, Girton College, Cambridge

2006 PhD Bristol


See publications page for full list and Google Scholar for list plus citations.


Stevens, M. Cheats and Deceits: How Animals and Plants Exploit and Mislead. 2016. Oxford University Press.

Stevens, M. Sensory Ecology, Behaviour, and Evolution. 2013. Oxford University Press.

Stevens, M. & Merilaita, S. (Editors). 2011. Animal Camouflage: From Mechanisms to Function. Cambridge University Press.

Journal Publications

Over 100 full journal publications in a range of journals spanning Nature, PNAS, Nature Communications, BMC Biology, Proceedings B, and many others.

Main and Recent External Grants

01/10/2017 – 30/09/2020: BBSRC Industrial Partnership Award (part funded by QinetiQ), £376,743. How to optimise imperfect camouflage.

01/12/2016 – 31/05/2017: BBSRC Pathfinder, £11,000. Imaging Animal Vision.

2017: Multiple Industry Grants, ca£40,000.

01/03/2017 – 31/03/2017: EasyFix, £3,175. Horse vision and fence design.

10/08/2016 – 24/08/2016: Fundação de Amparo à Pesquisa do Estado de São Paulo – FAPESP, £2,268. Visiting research position at University of São Paulo.

01/10/2014 – 30/09/2017: BBSRC, £371,695. Predator learning of camouflage types. (with Co-I John Skelhorn, Newcastle University).

01/08/2012 – 30/09/2014: BBSRC, £520,000. Predator Vision and Avian Egg Camouflage.

01/10/2009 – 03/03/2015: BBSRC David Phillips Fellowship: £1,002,266. Predator vision and defensive coloration: from mechanism to function.

Invited Lectures

Public Lectures

Royal Institution (London), Royal Geographical Society, Army and Navy Club (Washington DC), Hay-on-Wye Book Festival, Brighton Science Festival, Café Scientifique, Penzance Literary Festival, and others.

Academic Lectures

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).

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 (Genetics and Evolution), Switzerland (2015); Wiko Berlin, Animal Coloration Meeting.

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

Asia: National University of Singapore (Biology); International Primatological Conference, Kyoto, Japan (2010); Integrative Behavioral Biology, Xi’an, China (2011, Keynote); International Symposium on Avian Brood Parasitism, Hainan, China (2012); Camouflage Cultures: surveillance, communities, aesthetics, animals (2013), Sydney, Australia.

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

Media and Outreach

I have and currently work with a wide range of companies and organisations, including: Natural History Museum, London; California Academy of Sciences; Frieze Art Foundation; BBC; Rentokil Initial; QinetiQ; British Horseracing Authority; British Eventing; Greyhound Board of great Britain; EasyFix; Donkey Sanctuary; National Lobster Hatchery; Bird World; Zoological Lighting Institute.

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.

I am also currently a BBSRC Schools Regional Champion for the SW for my outreach work to school students.

Dr Jolyon Troscianko

BBSRC Post-Doctoral Research Associate

Centre for Ecology and Conservation, University of Exeter.

More About Jolyon

I have a background in behavioural ecology and sensory ecology, asking questions such as how an animal’s cognition or appearance affects how they interact with their environment, and how this in turn affects their evolution (see my publications below).

I’m currently a postdoctoral research associate, working on a BBSRC grant to Martin Stevens and John Skelhorn, investigating how easily predators can learn and switch between different camouflage strategies. I’m testing this using a series of touch-screen experiments on humans and laboratory chickens, training them to find computer generated “moths” that are concealed with different types of camouflage.

Combining my interests in photography, image analysis and animal vision has led me to develop new tools and methodologies for using standard consumer digital cameras as multispectral imaging tools for scientific purposes. This includes ultraviolet photography, and models that can convert from “camera vision” to “animal vision” (see the Image Analysis Toolbox).

My previous work included a postdoc with Martin and Claire Spottiswoode investigating how camouflage affects the survival of animals in the real world, using ground-nesting birds in Zambia as a model. We discovered that pattern and contrast matching were the most important cues that gave away the location of nests to predators, and that each female could choose where to create her nest based on the appearance of her own plumage, or the appearance of her clutch.

For my PhD I focussed on the ecological significance of tool-use in New Caledonian crows. Using methods ranging from animal-borne video cameras to isotope analysis and motion-triggered cameras, I contributed to our understanding of the importance of tool-use to the crows’ ecology and survival in this secretive species. For example, the isotope analysis suggested that over 1/3rd of the crows’ daily energy requirement could depend on tool-use, while animal-borne cameras revealed previously unknown foraging strategies and food sources. My research into the New Caledonian crows’ vision also highlighted how eye and bill morphology could have evolved to enable tool-use at the expense of other ecological pressures on their morphology such as accessing carrion.

Click here to see my publications


Emmanuelle Briolat

BBSRC-funded PhD student

Centre for Ecology and Conservation, University of Exeter.

More About Emmanuelle

Supervisors: Martin Stevens (Primary), Jon Blount

Working on the form and honesty of warning signals in insects.

Current Research

The aim of my PhD project is to investigate the form and function of aposematic signalling in insects, and in particular in Lepidoptera. I am especially interested in exploring the question of honesty in warningly coloured animals, or how the variation in the strength of their aposematic signals relates to the potency of their chemical or physical defences.

To address this problem, I am focusing on a charismatic family of day-flying moths, the burnet moths (Zygaenidae). They represent an ideal system in which to study both intra- and inter-specific variation in signals and defences: burnet moths sequester toxic cyanogenic glycosides from their host-plants and present a range of different colour patterns on their wings, varying both subtly within the Zygaeninae and more dramatically between subfamilies. To analyse signal honesty in the burnet moth system, I will be using a combination of photography, to measure colouration as perceived by potential predators, and toxicity analyses, using specimens collected from Cornwall, Denmark and a range of sites in France. I will also be performing fieldwork experiments with artificial prey, based on the observed variation in burnet moth signals, to tease apart the relative roles of different elements of colour patterns in protecting aposematic prey.

Previous Research

I graduated with a BA in Natural Sciences (Zoology) from the University of Cambridge in 2012. A summer project after my second year, working with Martin Stevens and Alexandra Torok on her startle display project, gave me my first taste of research in the field of sensory ecology. I then went on to work on two quite different short projects in my final year, first studying the relationship between the invasive zebra mussels and killer shrimp, then investigating the genes responsible for cyanogenesis in Heliconius butterflies.

After graduating, I returned to my interests in sensory ecology, studying cuttlefish camouflage patterns as part of a three month internship programme at the Marine Biological Laboratory in Woods Hole, MA. I then worked as a research assistant in Rebecca Kilner’s burying beetle group at Cambridge University, participating in fieldwork, laboratory experiments investigating the interactions of beetles with mites and microbes, and a long term selection experiment exploring the influence of body size on the beetle life cycle and behaviour.

Sara Mynott

NERC-funded PhD student

Centre for Ecology and Conservation, University of Exeter.

More About Sara

Supervisors: Martin Stevens (Primary), Steven Widdicombe (Plymouth Marine Laboratory)

Working on the impacts of climate change on intertidal species, camouflage and predation.

Current Research

In September 2014 I embarked on a PhD exploring how climate-related environmental change is affecting intertidal animals. I am particularly interested in how temperature and acidification influences the ability of crustaceans to camouflage, and, in turn, how this impacts their predation risk.

Camouflage is the most widespread way of avoiding predators, but stress can have a major impact on colour change in a number of species, and, as the climate changes, shifts in environmental conditions have the capacity to cause such stress.

I study camouflage in a variety of crustaceans, including common shore crabs, ghost crabs and chameleon prawns. By exposing them to different conditions and photographing them at regular intervals, we can find out how changing seawater properties affects their camouflage.

The photos can be used to model what individuals would look like from their predator’s perspective, and lets us discern whether one set of conditions leaves them more vulnerable to predation than another.

Previous Research

I completed my undergraduate in Environmental Geoscience at the University of Bristol in 2010 and moved into the marine realm, with a masters in Marine Ecology and Environmental Management at Queen Mary University of London in 2012.

My marine research to date has focussed on the temperature-size rule and its fitness trade-offs, as well as the methods used in crustacean fisheries management.

During my MSc, I explored the effect of temperature on the development and fitness of brine shrimp. By rearing the shrimp from cyst to adult, and measuring their size, survivorship and reproductive output, I was able to derive a possible mechanism for the temperature-size rule.

More recently, I conducted research into crustacean fisheries at University Marine Biological Station Millport (UMBSM). Field studies off the Scottish west coast coupled with aquarium studies at UMBSM allowed me to assess both how trap deployment and intraspecific interactions influenced total catch.

Victoria Lee

NERC-funded PhD student

Centre for Ecology and Conservation, University of Exeter.

More About Victoria


Alice Rosen

MSc by Research student

Centre for Ecology and Conservation, University of Exeter.

More About Alice

Supervisors: Martin Stevens (Primary), Andrew Young

Working on coincident disruptive coloration and background matching in the common frog.

Current research

The aim of my project is to investigate the anti-predator coloration strategies of the common frog. I will be exploring concealment through background matching, and I am particularly interested in how the markings and patterns of the common frog may be used to manipulate the receiver’s visual system in order to avoid detection.

The common frog, Rana temporaria, has a variety of mammalian and avian predators and exhibits extreme variation in coloration and pattern. It has been proposed that the common frog exhibits coincident disruptive coloration. This camouflage technique involves the use of markings arranged to create a false contour that is more conspicuous than the true body outline. In the common frog, barring on the hind legs, aligned in the resting position, is thought to break up the true boundary contours of the legs and merge distinct body regions, exploiting predator spatial vision through disrupting their perception of recognisable features.

Using artificial models of Rana temporaria in the field, as well as a computer-based human visual search experiment, I hope to figure out if a coincident disruptive boundary across the legs and body improve concealment, and also investigate whether this coloration has evolved to target the visual system of a particular predator.

I also aim to determine if common frogs are tuned to their background in terms of coloration and pattern. Using image analysis software to simulate the visual systems of common predators I will test whether frog color, luminance and pattern matches more closely to their own background than to backgrounds from other sites.

Previous research

I graduated from the University of Exeter in July 2016 with a First Class Honours degree in Zoology. For my undergraduate research project, supervised by Dr Chris Lowe, I investigated natural populations of the snakelocks anemone, which exists as a fluorescent green morph and a non-fluorescent brown morph. I examined the photosynthetic responses of the two morphs at a range of irradiances using photorespirometery to address the role of fluorescent proteins in light optimisation, in either a photo-protective or photo-enhancing capacity.

Previously, as Scientific Director of a student-led expedition team, I collected data on herpetological diversity in a remote peatland rainforest in Borneo over Summer 2015. We conducted a series of surveys, including visual encounter transects, pitfall traps and acoustic logging to evaluate the efficacy of the techniques and to aid conservation efforts by building a case for the protection of peatland rainforests.


Jenna Proctor

MSc by Research student

Centre for Ecology and Conservation, University of Exeter.

More About Jenna

Supervisors: Martin Stevens

Working on the adaptive camouflage of the furrowed crabs and investigating their ability to background match through moulting and behaviour.


Current research:

My project investigates how camouflage can be adapted to allow animals to move freely across environments that are highly variable. These gradual camouflage adaptations are becoming more commonly studied. Previous work by Martin Stevens has found that the European green shore crab (Carcinus maenas) are capable of using a mixture of behavioural colouration matching and consecutive moulting in order to match their carapace colouration to their surrounding environment. While many other intertidal animals are also thought to adopt this technique it has seldom been explored. I hope to use a mixture of field and laboratory experiments to investigate this strategy in the understudied furrowed crab (Xantho hydrophilus).

I plan to investigate how well the furrowed crab can visually camouflage their carapaces through background matching. I will set up tanks with various colours and degrees of brightness and allow crabs to moult while on these backgrounds. I will record their changes using digital camouflage and image analysis. This will allow me to assess the colour changes of each crab.

Previous research:

I graduated from the University of Exeter in July 2016 where I completed a BSc in Zoology. I carried out a final year project that focused on the trail following behaviour of the common garden snail (Cornu aspersum). I aimed to investigate whether the slime trail of the snail acted as an information channel for other snails. I kept the snails in varying conditions and assessed whether this impacted other snails’ decisions of whether to follow a particular slime trail or not.

In the summer of 2015 I undertook a short project at Cambridge University. This project was to review and compile existing literature and data to investigate male weaponry across all taxa.




Sam Green

MSc by Research student

Centre for Ecology and Conservation, University of Exeter.

More About Sam

Supervisors: Dr Martin Stevens (Primary), Dr Alastair Wilson

Working on the colour change and behavioural camouflage strategies used by the chameleon prawn (Hippolyte varians).

Current Research:

My research focuses on the chameleon prawn (Hippolyte varians), a species with a wide variety of coloured morphs, many of which subjectively show close resemblance to the seaweed that they are found on. Previous studies have shown that chameleon prawns can seemingly change colour over time to match novel algal substrates and that this colour change likely involves the expansion of pigments within specialised chromatophore cells.  Research has also shown that the species undergoes a circadian rhythm of colour change between a coloured day form and a blue ‘nocturne’. These characteristics could camouflage the chameleon prawn within the algal beds, potentially providing valuable protection against predators.

My aim is to investigate the behavioural and colour change strategies used for camouflage by chameleon prawns and how the combination of both strategies can give a competitive edge.  To investigate the behavioural elements my research will consist of determining natural distributions of coloured morphotypes and conducting choice experiments between algal substrates in the lab. For the physiological strategies we will be testing the degree to which colour change can occur in this species, and determining if this ability varies between morphotypes. I also plan to explore the potential link between diet and colour change using natural and artificial substrates. The analysis of colouration and luminance matching will be done using image analysis, through recording objective measures of reflectance and colour, then modelling colour change and camouflage to fish visual systems.

Additionally, among the various coloured morphotypes there exists a transparent morph. Compared to the other morphotypes, very little experimental work has been conducted on this morph and so there is scope for investigation to further understand the ecology and efficiency of camouflage of transparent prawns, and whether they represent a different strategy to the coloured individuals.

Previous Research:

I graduated from the University of Exeter in July 2016 (BSc Zoology 1st Class). My undergraduate research project was supervised by Professor Dave Hodgson investigating the use of fluorescence as a deterrent against avian predators by the common garden snail (Cornu aspersum).

I have interned with the Amphibian and Reptile Conservation Trust, gaining an accreditation under their licence to work with the sand lizard (Lacerta agalis), and as part of a research team in ZSL’s badger-cattle contact project in Cornwall. I have also worked as a research assistant in a behavioural study looking at the effects of female biased sex ratios on mating and fighting in flour beetles (Gnatocerus cornutus). During spring 2012 I spent time with the Department of Parks and Wildlife, The Gambia. I participated in biodiversity surveys and community engagement projects, and gained insight into conflicts between conservation aims and local communities.

I developed an interest in the field of sensory ecology after taking Martin’s module in the final year of my undergraduate degree and this has inspired me to conduct my research masters in this field. In my future research I will combine my interests in both conservation and sensory ecology to develop more effective conservation strategies and tools.





Roz Evans

Project Assistant

Centre for Ecology and Conservation, University of Exeter.

More About Roz

I’m a Project Assistant for the Sensory Ecology group, and my main aim is to increase and monitor the impact that the research group’s work could have in the outside world. With so many organisations working with animals, a lot of the research produced by the Sensory Ecology group can be useful, allowing them to understand how the animals they focus on see the world around them. This could be applied to increasing animal welfare, or making products better from the animal’s point of view.

Previous work

My professional work to date has been focused on science communication, largely through the production of Biosphere wildlife science magazine. I graduated 1st Class from Exeter University in Conservation Biology and Ecology a few years ago.  For my final year project, I investigated the effect of altitude on the size of blue tit eggs.





Are you interested in joining our group?

There are a number of opportunities in our lab, especially for work on camouflage in shore animals and other areas of anti-predator coloration. Please get in touch if you are interested.

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