Predator Vision and Avian Egg Camouflage

Lead Research Organisation: University of Exeter
Department Name: Biosciences

Abstract

Few things are more important to an animal than avoiding predation. As soon as Darwin and Wallace presented their theory of evolution by natural selection, around 150 years ago, the numerous ways of avoiding predation provided crucial examples for illustrating and defending natural selection and adaptation. Animal coloration has also influenced many areas of human applications and culture. For example, camouflage is of great importance in military applications and has influenced peaceful areas of culture such as art and fashion. Camouflage is one of the most important anti-predator adaptations in nature and crucial to the survival of individuals in countless species. It is a textbook example of natural selection.
Given the importance of camouflage, it is unsurprising that many different types exist. These include background matching, where the animal matches the general appearance of its environment, and disruptive coloration, where the outline of the animal's body is broken up by strongly contrasting markings. In addition, some animals may use masquerade, where they resemble an apparently unimportant object in the environment such as a leaf or stone. Within the last decade substantial research has investigated the different types of camouflage that may exist and how they may work in terms of predator vision. Studies of camouflage have been undertaken by evolutionary, sensory, molecular, and behavioural biologists, experimental psychologists, artists and computer scientists. Camouflage research is consequently a vibrant and interdisciplinary area and a model system for investigating animal behaviour, vision, and evolutionary biology. However, numerous gaps in our knowledge remain. Remarkably, given how important camouflage is, we still know little about it in real animals. While recent theoretical breakthroughs have been numerous, work has almost entirely been undertaken in artificial systems. Consequently, we know little of how such findings relate to the natural world. In short, camouflage theory from artificial systems greatly outstrips our understanding in real species and habitats. However, we can now use this theory to make a range of predictions about camouflage and its evolution in the wild.
In this project, we use camouflage in birds eggs as a model system to test several key questions about concealment, its value and evolution. Avian egg camouflage makes a wonderful system because the backgrounds on which many species nest is well known, substantial information about breeding biology exists, predators are often well documented, and bird eggs in many species have impressive camouflage. We will conduct research in South Africa and Zambia over two field seasons, using several groups of birds (mainly plovers and nightjars). We will locate nests of various species and undertake analyses of the colour and pattern of the eggs against the background on which they were laid. Using mathematical models of animal vision and techniques from computer science, we will analyse how camouflaged the different egg types are against the different available backgrounds, and determine the camouflage type that is present. In addition, we will check nests at regular intervals over the incubation period and determine which nests are eaten by predators and how long the different eggs survived. We will record predation events to determine the main predator groups for each species so that we can model egg appearance in terms of predator vision. Using these data, we will answer several key questions: 1) how does the level and type of camouflage affect the survival of an individual's eggs, 2) does the background structure influence the type of camouflage that evolves, and 3) how do species that nest on multiple background types optimise their level of concealment over evolution? Our study will reveal fundamental answers about how the type and level of camouflage influences survival in complex natural environments.

Technical Summary

Avoiding predation is a crucial aspect of many animals' fitness. A widespread defence is camouflage, which is a textbook example of natural selection. Animal camouflage has also influenced human applications and culture, including the military, art and fashion. In nature, many types of camouflage may exist. These include background matching, where the animal matches the appearance of its environment, disruptive coloration, where the body outline is broken up by contrasting markings, and masquerade, where animals resemble an unimportant object such as a leaf or stone. In the last decade a resurgent research agenda has investigated the types of camouflage that may exist and how they work in terms of predator vision. Studies of camouflage have been undertaken by evolutionary, sensory, and molecular biologists, psychologists, and computer scientists. However, we know little about camouflage in real animals because theoretical work has almost entirely been undertaken in artificial systems. Here, we use camouflage in bird eggs as a model system to test key questions about camouflage. Avian egg camouflage is a perfect system because the backgrounds on which species nest is well-known, and substantial information about breeding biology and predators exist. We will conduct research in South Africa and Zambia, using several groups of birds (mainly plovers and nightjars). We will locate nests and use computer models of animal vision and image analysis to quantify the level and type of camouflage present. In addition, we will obtain survival analysis of nests over time and record predation events to determine the main predator groups. Using these data, we will answer: 1) how the level and type of camouflage affects survival, 2) how background structure influences the type of camouflage that evolves, and 3) how species that nest on multiple backgrounds optimise concealment. Our study addresses fundamental issues about how camouflage influences survival in natural environments.

Planned Impact

(1) Public Outreach, Education and Learning
Animal coloration, including camouflage can readily attract significant media attention and captivate public interest. It is highly visual, and the concepts of predation and animal coloration are widely appealing and understandable. Such concepts generally receive coverage in popular science books, newspapers, television and radio, and exhibitions. Animal coloration and camouflage has inspired areas of art and fashion, including exhibitions at various galleries and museums. The PI has appeared on and helped with radio programmes and TV series, and his research has been covered on various occasions in the media, including National Geographic, New Scientist, the BBC, and the New York Times. He has dealt with journalists and press offices regularly including issuing press releases and interviews. As well as being ideal to appeal to both to the general public and media, the project also represents a key example of natural selection that can be used to illustrate a range of key concepts regarding evolution and natural history to children and young adults. It is a key aim of ours to communicate our work and its principles to these audiences. To this end, we will enrol as Science and Engineering Ambassadors to present our work at schools/colleges, and present displays and interactive encounters at the Cambridge Science Festival and the Royal Society Summer Science Exhibition. We will also set up a website containing high-quality images and videos and an educational game illustrating key principles of camouflage, predation, and evolution using visual aids from our fieldwork. We will also acquire a database of images and videos for use by the media, teachers, and artists. Finally, the digital imaging methods in our project would also allow the production of photographs showing how non-human animals (e.g. predators), 'see' the world around them. Our images and videos will also be valuable to go alongside press releases.
(2) Economic
Camouflage has long had significant application in the military, plus in human recreation and commerce (e.g. hunting and wildlife watching). The PI has organised a conference on animal vision and camouflage, which was attended by members of the military, and is cosupervising a CASE PhD project partly funded by the Ministry of Defence. In addition to designing camouflage, there is also a need for camouflage breaking. To do this, requires an understanding of how camouflage works in natural environments, and often utilises imaging approaches. There is also an increasing effort to disguise unattractive but functional human made objects, such as mobile phone transmission masts and satellite dishes, and successful ways of doing so would yield financial rewards. Once we have acquired and analysed sufficient data to have obtained preliminary findings, we will coordinate with relevant departments and organisations about the application and exploitation of our work. The key insights our project will give is information about what types of camouflage work best against different background and habitat types, and how camouflage can be optimised in complex real world systems (natural or anthropogenic).

Publications

10 25 50
 
Description The project has shown that several species of ground nesting bird select local microhabitats that better match the colour and pattern of their eggs compared to habitat patches further away.

Survival data have shown that high contrast eggs fare well on high contrast backgrounds, and low contrast eggs on low contrast backgrounds. However, adults that lay high contrast eggs on low contrast backgrounds are less likely to survive, and the predation risk is always high for eggs on a low contrast background.



Comparisons have shown that the egg colours and patterns of some species are matched to their backgrounds better than those of other species, and other nests from the same species.



Our citizen science egg and adult nightjar games have highlighted trade-offs between di- and trichromatic predator visual systems, with dichromats outperforming trichromats under certain circumstances where prey contrast and background contrast is high.
Exploitation Route Further research, outreach
Sectors Education

 
Description Imaging animal vision
Amount £10,842 (GBP)
Funding ID BB/P017339/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 12/2016 
End 05/2017
 
Description Predator learning of camouflage types Grant Ref: BB/L017709/1
Amount £371,695 (GBP)
Funding ID BB/L017709/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 10/2014 
End 09/2017
 
Description Collaboration with Estación Biológica de Doñana CSIC 
Organisation Spanish National Research Council (CSIC)
Department Doñana Biological Station
Country Spain 
Sector Public 
PI Contribution Collaboration with a PhD student on the camouflage of ground nesting birds at the Doñana Biological Station. We have exchanged ideas, trained him in some imaging techniques and provided equipment to help facilitate research.
Start Year 2014
 
Description Collaboration with a BBSRC David Phillips Research Fellow in the Department of Zoology at the University of Cambridge 
Organisation University of Cambridge
Country United Kingdom 
Sector Academic/University 
PI Contribution Collaboration with a BBSRC David Phillips Research Fellow in the Department of Zoology at the University of Cambridge has facilitated successful field work and data collection in Zambia and South Africa. Collaboration with Research Associate at the Percy FitzPatrick Institute of African Ornithology at the University of Cape Town.
Start Year 2012
 
Description Mr David Griffiths 
Organisation FoAM Kernow
Country United Kingdom 
Sector Private 
PI Contribution Collaboration an award winning game designer, creative coder and livecoding artist. Collaboration with David has resulted in the creation of three online citizen science games to investigate camouflage theories.
Start Year 2013
 
Title Multispectral Image Calibration and Analysis Toolbox 
Description Digital cameras can be powerful tools for measuring colours and patterns in a huge range of disciplines. However, in normal 'uncalibrated' digital photographs the pixel values do not scale linearly with the amount of light measured by the sensor. This means that pixel values cannot be reliably compared between different photos or even regions within the same photo unless the images are calibrated to be linear and have any lighting changes controlled for. Some scientists are aware of these issues, but lack convenient, user-friendly software to work with calibrated images, while many others continue to measure uncalibrated images. We have developed a toolbox that can calibrate images using many common consumer digital cameras, and for some cameras the images can be converted to "animal vision", to measure how the scene might look to non-humans. Many animals can see down into the ultraviolet (UV) spectrum, such as most insects, birds, reptiles, amphibians, some fish and some mammals, so it is important to measure UV when working with these animals. Our toolbox can combine photographs taken through multiple colour filters, for example allowing you to combine normal photographs with UV photographs and convert to animal vision across their whole range of sensitivities. The toolbox is free, open source and open access and works with the free programme Image J. It is linked to this paper: 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. 
Type Of Technology New/Improved Technique/Technology 
Year Produced 2015 
Impact Still early stage. Being used by wide range of scientists already. 
URL http://www.sensoryecology.com/image-analysis-tools/
 
Description BBC 1 TV Appearance 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact I appeared on BBC 1 'The One Show' about my work on camouflage and colour change in rockpool goby fish.

no actual impacts realised to date
Year(s) Of Engagement Activity 2014
 
Description BBC2 TV Appearance 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Media (as a channel to the public)
Results and Impact Appearance on a BBC 2 TV programme as part of a series called 'Inside the Animal Mind'

no actual impacts realised to date
Year(s) Of Engagement Activity 2014
 
Description California Academy of Sciences Exhibition 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Color of Life exhibition at the California Academy of Sciences in San Francisco, CA. I collaborated to modify our online citizen science bird camouflage games to make them part of this exhibition that will run for five years. Annual attendance is 1.4 million people and the public stated that the game is one of the most enjoyable aspects of the exhibition.
Year(s) Of Engagement Activity 2015,2016,2017
 
Description Color of Life exhibition at the California Academy of Sciences 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Color of Life exhibition at the California Academy of Sciences in San Francisco, CA. I collaborated to modify our online citizen science bird camouflage games to make them part of this exhibition that will run for five years. Annual attendance is 1.4 million people and the public stated that the game is one of the most enjoyable aspects of the exhibition.
Year(s) Of Engagement Activity 2015
 
Description Grand National Outreach Event 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Grand National at Aintree Racecourse. I am collaborating with Aintree Racecourse to make a visual display (a slideshow) of how the Grand National appears to horse vision. This will form part of a course walk and other events, and is likely to be seen by thousands of visitors each day.
Year(s) Of Engagement Activity 2017
 
Description Invited seminar series 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Geographic Reach International
Primary Audience Participants in your research or patient groups
Results and Impact Seminar series at University of California Davis, Santa Cruz, and LA, where the PI spoke about the project and early findings. Invited series of seminars by the PI at three UC universities in USA Oral presentations for each of UC Davis, UC LA, and UC Santa Cruz

no actual impacts realised to date
Year(s) Of Engagement Activity 2013
 
Description NHM Exhibition 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact NHM London 'Colour and Vision' exhibition. We collaborated with the NHM to make a new interactive citizen science game on camouflage (in shore crabs) and predator vision (played by over 6200 people), and to make a series of interactive image displays that show how a range of other animals see the world, from nautilus to dogs. The exhibition was visited by over 40,000 people and both exhibits were commented on by the public as some of the most engaging and enjoyable aspects of the exhibition.
Year(s) Of Engagement Activity 2016
 
Description NHM London 'Colour and Vision' exhibition 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact I collaborated with the NHM to make a new interactive citizen science game on camouflage (in shore crabs) and predator vision (played by over 6200 people), and to make a series of interactive image displays that show how a range of other animals see the world, from nautilus to dogs. The exhibition was visited by over 40,000 people and both exhibits were commented on by the public as some of the most engaging and enjoyable aspects of the exhibition.
Year(s) Of Engagement Activity 2016
 
Description Online Guardian Article 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Public/other audiences
Results and Impact GrrlScientist from the Guardian covers our work

No impact as of yet
Year(s) Of Engagement Activity 2014
URL http://www.theguardian.com/science/grrlscientist/2014/aug/27/eggcellent-citizen-science-evolution-of...
 
Description Press coverage - Daily Mail 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Public/other audiences
Results and Impact The Daily Mail produced an article on our research video with the BBSRC, promoting the citizen science games we created, field work and general science

no actual impacts realised to date
Year(s) Of Engagement Activity 2014
URL http://www.dailymail.co.uk/sciencetech/article-2565113/Can-YOU-spot-bird-incredible-video-reveals-on...
 
Description Press coverage - Telegraph 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Media coverage in the Telegraph of our work and video made with the BBSRC. The Telegraph produced an article on our work, promoting our citizen science games, field work and video created by the BBSRC.

no actual impacts realised to date
Year(s) Of Engagement Activity 2014
 
Description Press coverage of paper 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact Media coverage of paper:
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.
Covered by: BBC Cornwall (11 August 2015), Daily Mail (6 August 2015), the Independent, the Washington Post (7 August 2015), IFL Science (8 August 2015).
Year(s) Of Engagement Activity 2015
 
Description STEM Ambassador; PDRA 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach local
Primary Audience Schools
Results and Impact PDRA is a registered STEM ambassador PDRA is registered as a STEM ambassador; inspiring school children in STEM subjects (Science, Technology, Engineering and Mathematics).

no actual impacts realised to date
Year(s) Of Engagement Activity 2013
 
Description School Talk - Ibstock Place School London 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach local
Primary Audience Schools
Results and Impact PDRA provided a talk on vision and camouflage at a London school PDRA provided a talk on vision and camouflage at Ibstock Place school.

no actual impacts realised to date
Year(s) Of Engagement Activity 2013
 
Description School Talk - Mullion School 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Schools
Results and Impact RA was invited to Mullion school to provide an interactive session on camouflage for kids in years 8-9

No impact as of yet
Year(s) Of Engagement Activity 2014
 
Description Science in the square 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach local
Primary Audience Public/other audiences
Results and Impact Public outreach event in Falmouth as part of Exeter University event. We presented one of our 'find the nightjar' outreach games. Displayed an interactive exhibit introducing the science we conduct. Over 2500 people attended the event. Computer game.

no actual impacts realised to date
Year(s) Of Engagement Activity 2013
 
Description Science, is it for me? 2013 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Schools
Results and Impact An event to inspire school children to study science while redefining the stereotype of scientists in lab coats. We presented some of the work we do in our project.
60 students from several local schools were invited to the University of Exeter where scientists organised sessions for the pupils to experience and learn about the research being conducted at the university.



Our Group provided a series of activities including; an introduction to signalling and camouflage, a session on search images, discussion on startle displays and an activity to appreciate camouflage on different habitats.
Presentation and camouflage games illustrating research.

no actual impacts realised to date
Year(s) Of Engagement Activity 2013
 
Description TV appearance 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Media (as a channel to the public)
Results and Impact National Geographic television programme ('Jurassic CSI') - I appeared in one episode of the series discussing bird vision and coloration

no actual impacts realised to date
Year(s) Of Engagement Activity 2011
 
Description Where is that nest? 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact An online citizen science game.

no actual impacts realised to date
Year(s) Of Engagement Activity 2013,2014
URL http://nightjar.exeter.ac.uk/story/nest_game
 
Description Zambia - School outreach 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Schools
Results and Impact RA and PDRA provided an activity for school children to teach them some theories of camouflage. Conducted in both 2012 & 2013 field seasons. PDRA and RA provided a camouflage activity to teach school children about theories of camouflage, including an introductory talk and discussion at the end.

no actual impacts realised to date
Year(s) Of Engagement Activity 2012,2013
 
Description Zambia - University Seminar 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Participants in your research or patient groups
Results and Impact PDRA provided a seminar for the Copperbelt University in Zambia PDRA provided a seminar for students at the Copperbelt University in Zambia entitled 'Photography and image analysis in science' Oral presentation

no actual impacts realised to date
Year(s) Of Engagement Activity 2013