CURRENT ISSUES - PERSPECTIVES AND REVIEWSTribute to Tinber.docx

CURRENT ISSUES - PERSPECTIVES AND REVIEWS
Tribute to Tinbergen: Public Engagement in Ethology
Julie Hecht* & Caren B. Cooper†
* Doctoral Program in Psychology, The Graduate Center, City
University of New York, New York, NY, USA
† Cornell Lab of Ornithology, Ithaca, NY, USA
(Invited Review)
Correspondence
Julie Hecht, Department of Psychology,
Hunter College and The Graduate Center,
City University of New York, 695 Park
Avenue, New York, NY 10065, USA.
E-mail: [email protected]
Received: October 4, 2013
Initial acceptance: October 19, 2013
Final acceptance: November 25, 2013
(M. Hauber)
doi: 10.1111/eth.12199

Keywords: citizen science, data quality,
animal behavior, informal science education
Abstract
Public engagement in research, called citizen science, has led to
advances
in a range of fields like astronomy, ornithology, and public
health. While
volunteers have been making and sharing observations
according to pro-
tocols set by researchers in numerous disciplines, citizen
science practices
are less common in the field of animal behavior. We consider
how citizen
science might be used to address animal behavior questions at
Tinbergen’s
four levels of analysis. We briefly review resources and
methods for
addressing technical issues surrounding volunteer
participation—such as
data quality—so that citizen science can make long-standing
contributions
to the field of animal behavior.
Introduction

‘Citizen science’ describes the various ways that mem-
bers of the public participate in genuine scientific
research (Cooper et al. 2007a; Silvertown 2009; Shirk
et al. 2012). Many citizen science projects arise from
communities with specific concerns (such as long-
term environmental monitoring), and some are
initiated by scientists to address specific research
objectives. Other projects have additional goals to
increase science literacy (Bonney et al. 2009a).
Research via public engagement is possible because
people enjoy making natural history observations and
sharing these observations with professionals and
peers. In addition, Web 2.0 and mobile technologies
facilitate mass participation. Consequently, citizen
science projects have arisen in a myriad of disciplines,
including ecology, phenology, macroecology, public
health, natural resource management, hydrology,
urban planning, meteorology, math, volcanology, and

various taxon-specific fields such as entomology,
ornithology, and herpetology. In recent years, citizen
science has enabled substantial contributions to the
fields of astronomy (Lintott et al. 2008), medicine
(Khatib et al. 2011a,b), and climate change (Morisette
et al. 2009).
Currently, the field of animal behavior is under-
represented among citizen science projects. Animal
behavior research integrates diverse methodological
approaches to investigate a wide array of scientific
questions about the behavior of wild and domesti-
cated animals in natural and captive settings. Public
observations of occurrence data (e.g., wildlife sight-
ings; Sn€all et al. 2011), count data (e.g., number of
birds; Cooper et al. 2007b), simple measurements
(e.g., rainfall; Cifelli et al. 2005), and even phenologi-
cal stages (e.g., budburst; Morisette et al. 2009) are
more common applications of citizen science, in part

because such data likely require simpler protocols
than most animal behavior data.
The dearth of citizen science research in animal
behavior is not necessarily due to lack of general pub-
lic interest. Pet-keeping is a global phenomenon
(Serpell 1996); wildlife tourism and wildlife docu-
mentaries are popular (Reynolds & Braithwaite 2001;
Nelson & Fijn 2013); zoo attendance is on the rise in
parts of the world (Davey 2007); and people are
already expressing interest in animal behavior citizen
Ethology 120 (2014) 207–214 © 2013 Blackwell Verlag GmbH
207
Ethology
ethology
international journal of behavioural biology
science projects (Foster et al. 2011; Williams et al.
2012). In the USA, one in four adults watches wildlife
(USFWS 2011), and the numerous websites that

stream live video of pets (e.g., SPCA cams) and wild-
life (e.g., nesting birds) are extremely popular (e.g.,
the Cornell Lab of Ornithology nest cams receive
millions of unique viewers from 175 countries,
including up to 9,000 simultaneous views of live-
streaming video).
Citizen science contributions could enhance animal
behavior investigations by increasing observations on
time and geographic scales (e.g., latitudinal gradients,
urbanization gradients, comparing populations, etc),
and these projects also have the potential to process
massive amounts of data (e.g., video or photo tag-
ging). In essence, citizen scientists provide an extra
pair of hands—or extra thousands of pairs. While a
small team of researchers documented the behavior of
the colonial orb-weaving spider during a solar eclipse
in Veracruz, Mexico (Uetz et al. 1994), the team
might have used citizen scientists to capture behav-
ioral data on additional populations or species during

the short-lived event. Without the help of citizen sci-
entists, researchers are effectively ‘missing’ variation
among animals.
Field research, behavior observation, and behavior
coding can be time-consuming, challenging to coordi-
nate, and expensive. Trained volunteers might offer
lower-cost options for collecting and/or scoring
behavior (Williams et al. 2012). Depending on the
study design and complexity, volunteers can be used
to ensure blind coding.
Citizen science methods could facilitate access to a
wide variety of species in different contexts and foster
integration of studies of natural populations and pop-
ulations under artificial selection. Additionally, an
expanded research platform offers the possibility of
capturing animals in novel contexts. Video and social
media, particularly YouTube, are rife with examples
of uncommon (Burn 2011) or previously unexplored

behaviors, such as interspecies play or object play
(Nelson & Fijn 2013). Behaviors captured in real-
world contexts can lead to new research directions,
and citizen scientists could provide behavioral video
footage.
We examine the realized and potential application
of citizen science to research animal behavior—the
inquiry about what animals do, how they do it, and
why they do it. Tinbergen (1963) framed the disci-
pline as encompassing four levels of analysis: causa-
tion, development, function, and phylogeny.
We use Tinbergen’s levels of analysis as a frame-
work to discuss a possible partnership between citizen
science and animal behavior research. We note that
citizen scientists could examine a particular behavior,
for example bird song, to generate data for answering
questions at all four levels (e.g., causation: ‘What day
length triggers bird song?’; development: ‘Do offspring

imitate songs of parents?’; function: ‘Do females
prefer certain song types?’; and phylogeny: ‘Which
species have similar songs?’). Professional scientists
can obtain a more robust understanding of an
observed behavior through investigations of non-
competing hypotheses at multiple levels. Other
questions might be best suited for investigation at one
particular level of analysis. We discuss a variety of
applications below, with the common theme that citi-
zen science participants can scale-up and advance
research that individual scientists could not do alone.
Additionally, we review the technical facets of a
union between animal behavior and citizen science,
giving attention to crucial details in training volun-
teers and handling volunteer data.
Citizen Science at Four Levels of Analysis
In the 1963 paper, On aims and methods of Ethology,
Tinbergen proposed an integrated approach to the

study of ethology to address the lack of a unified
public image of ethology and to resolve conceptual
differences between researchers. Since then, “how
questions,” exploring the immediate causes of behav-
ior, and “why questions,” exploring the evolutionary
forces behind behavior, have been pursued as
complementary approaches.
Causation
The causal basis of behavior focuses on stimuli and
mechanisms that trigger, cue, or precede a behavior.
For early ethologists, physiological and neural mecha-
nisms occurred in a ‘black box’ upon which they
could only speculate. Today, inquiry into causation
draws from genetics, endocrinology, physiology, and
neurobiology. Direct participation and input from citizen
scientists is already underway in neurobiology (Eyewire,
https://eyewire.org/signup) and genetics (Phylo) (Coo-
per et al. 2013). For domesticated and wild populations,

the collection of saliva and fecal samples could contrib-
ute to studies of stress physiology and measurements of
well-being to ultimately address a wide variety of behav-
ioral endocrinological questions.
Participant involvement could complement labo-
ratory findings by collecting behavioral data on
pre-screened or pre-selected populations. For exam-
ple, volunteers might assist research into relationships
Ethology 120 (2014) 207–214 © 2013 Blackwell Verlag
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Public Engagement in Ethology J. Hecht & C. B. Cooper
between genetic markers and behavior by collecting
behavioral data on pre-selected animals (e.g., through
a web portal for annotating video) whose genes have
been mapped. Researchers recently determined that
particular allelic qualities of DRD4 and TH gene poly-
morphisms were associated with activity–impulsivity
in German Shepherds and Siberian Huskies, with

behavioral analysis stemming from approx. 250 dogs
(Kubinyi et al. 2012; Wan et al. 2013). Continued
exploration of gene variants in dogs could be coupled
with citizen science projects collecting behavioral data
on dogs with known genetic composition to generate
a more comprehensive understanding of behavioral
traits. Citizen science allows researchers to expand the
size of their data set without requiring the participants
to have any knowledge of the DNA sampling and gen-
otyping that go into pre-screening decisions.
The addition of a citizen science component can
increase the rigor of the research in at least two ways.
First, the volunteer contributions could ensure a dou-
ble-blind experiment. Second, the volunteers provide
observations from multiple observers, and researchers
can use a consensus tool to score behavior, analogous
to consensus tools (replication by multiple partici-
pants) common to online crowdsourcing efforts for

other research (Wiggins et al. 2011). Additionally,
volunteers can install cameras to record animal
behavior and facilitate remote coding by professional
researchers. For example, scientists used footage from
volunteer nest cameras to examine proximate control
of egg-laying behaviors in nestbox-dwelling bird
species (Cooper et al. 2009).
Development
Approaches to animal behavior from a developmental
perspective are also mechanistic, but with a focus on
changes in an organism’s capacity for expressing or
acquiring behaviors at different stages over the course
of a lifetime.
Citizen science offers a unique opportunity to inves-
tigate ontogenetic questions in a wide range of
species. For example, citizen scientists might assist in
the study of social behavior and document the devel-
opment of particular behavioral patterns—such as fac-
tors affecting the onset and development of play in

various species—or personality changes (Stamps &
Groothuis 2010). Volunteers could collect and report
data on single organisms—whether subjects be a com-
panion animal, a backyard nestbox or an annually
returning species—and generate substantial data
points for each individual. Ultimately, with many
volunteers, the increased sample sizes could make
full-scale statistical analysis possible (Dickinson et al.
2010).
Citizen scientists could also examine the develop-
ment of animal sensorium, such as in relation to mor-
phological traits. For example, assumptions about the
different sensory abilities in dog breeds abound, and
such differences might be expected given physiologi-
cal differences (McGreevy et al. 2003), but actual
behavioral and development data are wanting.
Researchers could design and implement a project
where participants conduct simple, in-home tests
examining behavior responses toward different stim-

uli at different ages, thereby offering additional data
on breed (and even species) sensory development.
These behavioral measures of development are com-
parable to traditional laboratory experiments (Lord
2013). But, with the help of citizen scientists,
researchers can obtain data on a much larger scale.
Zoo visitors and wildlife tourists can play a role in
monitoring individuals’ development. Zoo visitors
express interest in animal behavior studies (Bowler
et al. 2012) and could collect many more hours (or
years) of observations than possible by resource-
limited staff with many other care-taking duties. Such
efforts might elucidate the onset or development of
behaviors such as stereotypic motor patterns as well as
factors affecting the development of these behaviors.
Zoo visitors, however, can have bias toward collecting
observations of active animals (Altman 1998; Williams
et al. 2012).

Function
The analysis of function focuses on the pressures that
shape behaviors. The pervasive influence of the built
environment means that many organisms reside in
environmental conditions that differ in novel ways
from the conditions under which traits evolved. For
example, the anthropogenic sound levels near high-
ways and flight paths create novel selection pressures,
and some animals might be able to adjust their behav-
iors within a range of plasticity, such as birds singing
at higher pitches near roads (Slabbekoorn & Peet
2003; Slabbekoorn & den Boer-Visser 2006). Other
traits might exhibit limited phenotypic plasticity, and
disturbances may result in population decline. Fur-
thermore, both domesticated and non-domesticated
animals residing in homes or zoos will demonstrate
behaviors that might no longer be adaptive. Research-
ers can design participatory projects in which volun-

teers collect data on species in different environments
and investigate different hypotheses behind complex
behaviors. In an online project called CamClickr, par-
209
J. Hecht & C. B. Cooper Public Engagement in Ethology
ticipants added behavior-category tags to images
archived from nest cameras to help investigate varia-
tions in nest attendance (Voss & Cooper 2010).
Because funding cycles offered by granting institu-
tions tend to span three- to five-year periods, collect-
ing long-term data sets can be challenging for
research scientists. The collection of such data sets
could be made possible through citizen science activi-
ties. Depending on the initiative, initial investment to
set up the project, and maintenance costs, could be
low with free online tools for managing projects (e.g.,
www.citsci.org), recruiting volunteers (e.g., www.

scistarter.com), and data archiving (e.g., www.data-
one.org). Citizen scientists could measure different
components of fitness such as survival to maturity,
longevity, mating success, and reproductive success.
Phylogeny
Behaviors can also be understood from a phylogenetic
perspective. By considering ancestry, researchers can
identify traits that are highly conserved and those that
are more recently evolved within the same lineage.
Because closely related species are found in everyday
environments, a citizen science project could collect a
wide variety of behavioral data on closely related spe-
cies in different environments that have adopted dif-
ferent survival strategies. By comparing closely
related species occurring on different continents (e.g.,
black-billed bagpie Pica hudsonia in America and the
European magpie Pica pica in Eurasia), citizen scien-
tists can provide simultaneous data sets without the

confound of temporal displacement typical of smaller-
scale research projects.
Not all citizen engagement in animal behavior pro-
jects fits neatly into Tinbergen’s levels of analysis.
Recently, Project: Play With Your Dog (Horowitz Dog
Cognition Lab, Barnard College) requested short vid-
eos depicting play between a dog and a person, with
the aim of capturing the wide array of behaviors and
conditions that encompass interspecific play. Video
submissions drew from around the world, and
although participants did not contribute directly to
behavioral analysis, citizen scientists had to think
about behavior in terms of how they and their dog
play together and provide behavioral descriptors in an
accompanying questionnaire.
Similarly, an online survey by Animal Welfare Indi-
cators asked participants to assist in developing a con-
tinuous behavioral scale of lameness for goats.

Participants watched nine videos and scored abnormal
gait, head-nodding, and spine curvature on a visual
analogue scale. Before participants began, each
behavior had a clear description, and participants
were schooled in measuring behavior as per the speci-
fications of the project.
Merging Ethology and Citizen Science Methods
Creating Successful Citizen Science Animal Behavior
Projects
Like all successful research endeavors, the design of
citizen science projects follows the steps of the scien-
tific method; the only difference is the additional layer
of complexity introduced by the involvement of the
lay public. Projects start with the formulation of scien-
tific questions, organization of a team of skilled
experts in relevant areas, and determination of appro-
priate data collection (Bonney et al. 2009b). Depend-
ing on the legal status and various protections offered

to different species, researchers might need to present
protocols to Institutional Animal Care and Use
Committees (Bayne 1998). Citizen scientists can learn
behavior-coding techniques—or relevant project-
related skills—in online videos or in person. The
research team can control quality before the project
begins by testing each prospective participant’s
accuracy and ability to collect meaningful data.
Behavioral data accuracy is a key concern, and pro-
jects incorporating citizen science data should con-
sider topics like observation quality, sampling bias,
and interobserver reliability (Dickinson et al. 2010;
Burghardt et al. 2012; Hart et al. 2012). Williams
et al. (2012) found that although zoo visitor interest
in the project was high, the activity budget data col-
lected on a group of captive otters were not accurate,
mostly because visitors did not adhere to the observa-
tion period; the researchers suggest that certain
sampling methods might be easier—or harder—for

observers.
Meanwhile, there are numerous ways to improve
data quality in citizen science (Cohn 2008; Silvertown
2009; Wiggins et al. 2011; Bonter & Cooper 2012).
Technologies like smartphones, multimedia, or inter-
active video could assist in time-keeping and behav-
ioral data collection (Parr et al. 2004; Aanensen et al.
2009). Data management tools, often through cyber-
infrastructure, improve efficiency (Newman et al.
2011). Additionally, some citizen science projects
demonstrate that certain types of people collect more
accurate data than others (Delaney et al. 2008), and
similar lessons can make their way into citizen science
projects in animal behavior.
Finally, researchers can be creative in where and
how volunteers access animals and how behavior will
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be observed or documented. In some cases, behaviors
are not observed directly but are inferred. Phenology
research addresses the timing of life-cycle events,
including behaviors such as egg laying and migration
in birds, insects, and some reptiles and amphibians
(Head et al. 2013). Limited access to animals in situa-
tions where volunteers can obtain unbiased observa-
tions of life-cycle events may have been one of the
largest impediments to citizen science in animal
behavior. Now, Internet and remote-camera techno-
logies combined with crowdsourcing infrastructure
offer new opportunities to collect data on animals that
are difficult to access (Deng et al. 2012; Head et al.
2013; Loos & Ernst 2013). Live-streaming cameras
can be set up at areas with known activity—animal
nesting or gathering sites—so that people can docu-
ment behaviors at a distance. In addition to remote/
web-mediated studies, possibilities abound for studies

of companion animals (e.g., dogs, cats, rabbits, birds,
ferrets, fish, gerbils, and hamsters), zoo animals, ani-
mals viewed during nature tourism, and urban wild-
life such as squirrels, pigeons, rats, foxes, and coyotes.
Volunteers could document the behavior of animals
at zoos or wildlife tourism destinations, especially ani-
mals that handlers and keepers do not have time to
record. Special interest groups—such as the House
Rabbit Society, backyard chicken groups, and pet
breeders—offer significant potential for citizen science
projects.
Training and Resources for Animal Behavior Inquiry
Skill in observing and measuring animal behavior
does not necessarily follow from an appreciation of or
even cohabitation with animals. Members of the pub-
lic are generally interested in animals yet have a pro-
clivity not toward Tinbergen’s four levels of behavior,
but often toward anthropomorphic explanations of
behavior.

While the study of animal behavior is an attempt to
investigate ultimate and proximate questions about
behavior, humans often characterize animal behavior
through an anthropocentric lens (Foster et al. 2011).
People readily ascribe human characteristics to non-
humans. Subjective assessments of non-human
behavior can be consistent across observers, often
reifying existing beliefs regardless of whether they
have been substantiated (Bahlig-Pieren & Turner
1999; Morris et al. 2008; Horowitz 2011). Seminal
work by Heider & Simmel (1944) found that people
consistently attribute human emotions, aims, and
objectives to inanimate objects moving in a ‘human-
like’ manner. Companion animal species, mammals,
and non-human primates are more readily anthropo-
morphized than other species (Eddy et al. 1993), even
though seemingly ‘alien’ species, like cephalopods,
display complex behaviors and cognitive processes

(Fiorito & Scotto 1992; Moriyama & Gunji 1997).
Importantly, the appearance of behavioral homologies
should not imply similar meaning, such as the ‘smil-
ing’ chimpanzee or dolphin (Horowitz 2007).
Tendencies toward anthropomorphism are perhaps
a useful ‘hook’ when engaging the general public
in animal behavior projects. Recently, an online
video titled ‘Lol cats like stroking too’ (http://www.
youtube.com/watch?v=OFCRvjle2o8) accompanied a
publication on the neurons activated by massage-like
stroking to assist in public understanding of the
research (Vrontou et al. 2013). The video received
over 200,000 views in the first year.
Clearly defined project expectations and training
materials can minimize the effects of anthropomor-
phic inferences (Voss & Cooper 2010). Scientists can
provide a detailed ethogram, an explicit coding
scheme, and a clear behavior-sampling technique

(Martin & Bateson 2007). Behaviors in an ethogram
can be distinguished based on structural or empirical
characteristics, such as ‘raised lips at edges’, or conse-
quence-based descriptors, such as ‘threat’. Empirical
descriptors help create unambiguous behavioral
parameters (Lehner 1987) but might offer challenges
for unpracticed observers. Volunteers need a clear
understanding of the terms in which they are
expected to view and document behavior and have
the technology, such as a timer, to code appropriately.
Researchers determine who is being watched and
when and how to watch them. Some coding schemes
provide information on how much time an animal
spends performing various behaviors, while other
schemes monitor behavior frequencies (Altmann
1974). Researchers must match the skill level of
observers with the scientific question. The role of vol-
unteers in the study of animal behavior will largely

involve the production of meaningful behavioral data.
The challenge will be to offer effective guidance and
instruction to avoid anthropocentric assessments.
A number of institutions have created behavior-
observation instructional platforms. Ethosearch (www.
ethosearch.org), created by the Lincoln Park Zoo, is a
resource for researchers and students to store and view
ethograms and practice measuring animal behavior.
Living Links (www.living-links.org)—a joint enclosure
for squirrel monkeys (Saimiri sciureus) and capuchins
(Sapajus apella) at the Edinburgh Zoo—offers a
public-learning resource to observe and score the behav-
ior of both species (Macdonald & Whiten 2011). Cam-
211
Clickr was an online citizen science project to code the
behaviors of nesting birds from still images as a way of

carrying out scan sampling. Voss & Cooper (2010) cre-
ated a college-level laboratory unit in which students
can use CamClickr to create ethograms while learning
about observer bias, observation techniques, and defin-
ing and quantifying observations.
Conclusion
Animal behavior researchers might consider whether
and how citizen science could support and amplify
their studies. Volunteers can expand the scope and
scale of animal behavior studies by bringing attention
to novel or rare behaviors or providing access to
additional variation in animal behaviors. Volunteers,
too, can benefit from personal experiences engaging
in hands-on scientific interactions with animals.
Undoubtedly, challenges arise with the involve-
ment of citizen science in animal behavior, yet the
possibilities are great. Altmann’s (1974, p. 229) text
on behavior sampling methods stresses that there

should not be a disconnect between the laboratory
and the real world: ‘Such a restriction on research
would mean that the behavioral sciences would for-
ever forsake any hope of knowing whether their most
powerful theories have any relevance in the world of
behavior outside the laboratory.’ Citizen scientists
offer an additional, and untapped, opportunity for
ethological exploration.
Acknowledgements
Thanks to Felicity Muth, Mark Hauber, and Andrew
Fulmer for conceptual conversations. Many thanks to
reviewers for comments, which improved the manu-
script.
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CURRENT ISSUES - PERSPECTIVES AND REVIEWS
Tribute to Tinbergen: Public Engagement in Ethology
Julie Hecht* & Caren B. Cooper†
* Doctoral Program in Psychology, The Graduate Center, City
University of New York, New York, NY, USA
† Cornell Lab of Ornithology, Ithaca, NY, USA
(Invited Review)
Correspondence

Julie Hecht, Department of Psychology,
Hunter College and The Graduate Center,
City University of New York, 695 Park
Avenue, New York, NY 10065, USA.
E-mail: [email protected]
Received: October 4, 2013
Initial acceptance: October 19, 2013
Final acceptance: November 25, 2013
(M. Hauber)
doi: 10.1111/eth.12199
Keywords: citizen science, data quality,
animal behavior, informal science education
Abstract
Public engagement in research, called citizen science, has led to
advances
in a range of fields like astronomy, ornithology, and public
health. While
volunteers have been making and sharing observations
according to pro-
tocols set by researchers in numerous disciplines, citizen
science practices

are less common in the field of animal behavior. We consider
how citizen
science might be used to address animal behavior questions at
Tinbergen’s
four levels of analysis. We briefly review resources and
methods for
addressing technical issues surrounding volunteer
participation—such as
data quality—so that citizen science can make long-standing
contributions
to the field of animal behavior.
Introduction
‘Citizen science’ describes the various ways that mem-
bers of the public participate in genuine scientific
research (Cooper et al. 2007a; Silvertown 2009; Shirk
et al. 2012). Many citizen science projects arise from
communities with specific concerns (such as long-
term environmental monitoring), and some are
initiated by scientists to address specific research
objectives. Other projects have additional goals to
increase science literacy (Bonney et al. 2009a).

Research via public engagement is possible because
people enjoy making natural history observations and
sharing these observations with professionals and
peers. In addition, Web 2.0 and mobile technologies
facilitate mass participation. Consequently, citizen
science projects have arisen in a myriad of disciplines,
including ecology, phenology, macroecology, public
health, natural resource management, hydrology,
urban planning, meteorology, math, volcanology, and
various taxon-specific fields such as entomology,
ornithology, and herpetology. In recent years, citizen
science has enabled substantial contributions to the
fields of astronomy (Lintott et al. 2008), medicine
(Khatib et al. 2011a,b), and climate change (Morisette
et al. 2009).
Currently, the field of animal behavior is under-
represented among citizen science projects. Animal
behavior research integrates diverse methodological

approaches to investigate a wide array of scientific
questions about the behavior of wild and domesti-
cated animals in natural and captive settings. Public
observations of occurrence data (e.g., wildlife sight-
ings; Sn€all et al. 2011), count data (e.g., number of
birds; Cooper et al. 2007b), simple measurements
(e.g., rainfall; Cifelli et al. 2005), and even phenologi-
cal stages (e.g., budburst; Morisette et al. 2009) are
more common applications of citizen science, in part
because such data likely require simpler protocols
than most animal behavior data.
The dearth of citizen science research in animal
behavior is not necessarily due to lack of general pub-
lic interest. Pet-keeping is a global phenomenon
(Serpell 1996); wildlife tourism and wildlife docu-
mentaries are popular (Reynolds & Braithwaite 2001;
Nelson & Fijn 2013); zoo attendance is on the rise in
parts of the world (Davey 2007); and people are

already expressing interest in animal behavior citizen
207
Ethology
ethology
international journal of behavioural biology
science projects (Foster et al. 2011; Williams et al.
2012). In the USA, one in four adults watches wildlife
(USFWS 2011), and the numerous websites that
stream live video of pets (e.g., SPCA cams) and wild-
life (e.g., nesting birds) are extremely popular (e.g.,
the Cornell Lab of Ornithology nest cams receive
millions of unique viewers from 175 countries,
including up to 9,000 simultaneous views of live-
streaming video).
Citizen science contributions could enhance animal
behavior investigations by increasing observations on
time and geographic scales (e.g., latitudinal gradients,

urbanization gradients, comparing populations, etc),
and these projects also have the potential to process
massive amounts of data (e.g., video or photo tag-
ging). In essence, citizen scientists provide an extra
pair of hands—or extra thousands of pairs. While a
small team of researchers documented the behavior of
the colonial orb-weaving spider during a solar eclipse
in Veracruz, Mexico (Uetz et al. 1994), the team
might have used citizen scientists to capture behav-
ioral data on additional populations or species during
the short-lived event. Without the help of citizen sci-
entists, researchers are effectively ‘missing’ variation
among animals.
Field research, behavior observation, and behavior
coding can be time-consuming, challenging to coordi-
nate, and expensive. Trained volunteers might offer
lower-cost options for collecting and/or scoring
behavior (Williams et al. 2012). Depending on the
study design and complexity, volunteers can be used

to ensure blind coding.
Citizen science methods could facilitate access to a
wide variety of species in different contexts and foster
integration of studies of natural populations and pop-
ulations under artificial selection. Additionally, an
expanded research platform offers the possibility of
capturing animals in novel contexts. Video and social
media, particularly YouTube, are rife with examples
of uncommon (Burn 2011) or previously unexplored
behaviors, such as interspecies play or object play
(Nelson & Fijn 2013). Behaviors captured in real-
world contexts can lead to new research directions,
and citizen scientists could provide behavioral video
footage.
We examine the realized and potential application
of citizen science to research animal behavior—the
inquiry about what animals do, how they do it, and
why they do it. Tinbergen (1963) framed the disci-

pline as encompassing four levels of analysis: causa-
tion, development, function, and phylogeny.
We use Tinbergen’s levels of analysis as a frame-
work to discuss a possible partnership between citizen
science and animal behavior research. We note that
citizen scientists could examine a particular behavior,
for example bird song, to generate data for answering
questions at all four levels (e.g., causation: ‘What day
length triggers bird song?’; development: ‘Do offspring
imitate songs of parents?’; function: ‘Do females
prefer certain song types?’; and phylogeny: ‘Which
species have similar songs?’). Professional scientists
can obtain a more robust understanding of an
observed behavior through investigations of non-
competing hypotheses at multiple levels. Other
questions might be best suited for investigation at one
particular level of analysis. We discuss a variety of
applications below, with the common theme that citi-

zen science participants can scale-up and advance
research that individual scientists could not do alone.
Additionally, we review the technical facets of a
union between animal behavior and citizen science,
giving attention to crucial details in training volun-
teers and handling volunteer data.
Citizen Science at Four Levels of Analysis
In the 1963 paper, On aims and methods of Ethology,
Tinbergen proposed an integrated approach to the
study of ethology to address the lack of a unified
public image of ethology and to resolve conceptual
differences between researchers. Since then, “how
questions,” exploring the immediate causes of behav-
ior, and “why questions,” exploring the evolutionary
forces behind behavior, have been pursued as
complementary approaches.
Causation
The causal basis of behavior focuses on stimuli and

mechanisms that trigger, cue, or precede a behavior.
For early ethologists, physiological and neural mecha-
nisms occurred in a ‘black box’ upon which they
could only speculate. Today, inquiry into causation
draws from genetics, endocrinology, physiology, and
neurobiology. Direct participation and input from citizen
scientists is already underway in neurobiology (Eyewire,
https://eyewire.org/signup) and genetics (Phylo) (Coo-
per et al. 2013). For domesticated and wild populations,
the collection of saliva and fecal samples could contrib-
ute to studies of stress physiology and measurements of
well-being to ultimately address a wide variety of behav-
ioral endocrinological questions.
Participant involvement could complement labo-
ratory findings by collecting behavioral data on
pre-screened or pre-selected populations. For exam-
ple, volunteers might assist research into relationships
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between genetic markers and behavior by collecting
behavioral data on pre-selected animals (e.g., through
a web portal for annotating video) whose genes have
been mapped. Researchers recently determined that
particular allelic qualities of DRD4 and TH gene poly-
morphisms were associated with activity–impulsivity
in German Shepherds and Siberian Huskies, with
behavioral analysis stemming from approx. 250 dogs
(Kubinyi et al. 2012; Wan et al. 2013). Continued
exploration of gene variants in dogs could be coupled
with citizen science projects collecting behavioral data
on dogs with known genetic composition to generate
a more comprehensive understanding of behavioral
traits. Citizen science allows researchers to expand the
size of their data set without requiring the participants
to have any knowledge of the DNA sampling and gen-

otyping that go into pre-screening decisions.
The addition of a citizen science component can
increase the rigor of the research in at least two ways.
First, the volunteer contributions could ensure a dou-
ble-blind experiment. Second, the volunteers provide
observations from multiple observers, and researchers
can use a consensus tool to score behavior, analogous
to consensus tools (replication by multiple partici-
pants) common to online crowdsourcing efforts for
other research (Wiggins et al. 2011). Additionally,
volunteers can install cameras to record animal
behavior and facilitate remote coding by professional
researchers. For example, scientists used footage from
volunteer nest cameras to examine proximate control
of egg-laying behaviors in nestbox-dwelling bird
species (Cooper et al. 2009).
Development
Approaches to animal behavior from a developmental

perspective are also mechanistic, but with a focus on
changes in an organism’s capacity for expressing or
acquiring behaviors at different stages over the course
of a lifetime.
Citizen science offers a unique opportunity to inves-
tigate ontogenetic questions in a wide range of
species. For example, citizen scientists might assist in
the study of social behavior and document the devel-
opment of particular behavioral patterns—such as fac-
tors affecting the onset and development of play in
various species—or personality changes (Stamps &
Groothuis 2010). Volunteers could collect and report
data on single organisms—whether subjects be a com-
panion animal, a backyard nestbox or an annually
returning species—and generate substantial data
points for each individual. Ultimately, with many
volunteers, the increased sample sizes could make
full-scale statistical analysis possible (Dickinson et al.
2010).
Citizen scientists could also examine the develop-

ment of animal sensorium, such as in relation to mor-
phological traits. For example, assumptions about the
different sensory abilities in dog breeds abound, and
such differences might be expected given physiologi-
cal differences (McGreevy et al. 2003), but actual
behavioral and development data are wanting.
Researchers could design and implement a project
where participants conduct simple, in-home tests
examining behavior responses toward different stim-
uli at different ages, thereby offering additional data
on breed (and even species) sensory development.
These behavioral measures of development are com-
parable to traditional laboratory experiments (Lord
2013). But, with the help of citizen scientists,
researchers can obtain data on a much larger scale.
Zoo visitors and wildlife tourists can play a role in
monitoring individuals’ development. Zoo visitors
express interest in animal behavior studies (Bowler

et al. 2012) and could collect many more hours (or
years) of observations than possible by resource-
limited staff with many other care-taking duties. Such
efforts might elucidate the onset or development of
behaviors such as stereotypic motor patterns as well as
factors affecting the development of these behaviors.
Zoo visitors, however, can have bias toward collecting
observations of active animals (Altman 1998; Williams
et al. 2012).
Function
The analysis of function focuses on the pressures that
shape behaviors. The pervasive influence of the built
environment means that many organisms reside in
environmental conditions that differ in novel ways
from the conditions under which traits evolved. For
example, the anthropogenic sound levels near high-
ways and flight paths create novel selection pressures,
and some animals might be able to adjust their behav-

iors within a range of plasticity, such as birds singing
at higher pitches near roads (Slabbekoorn & Peet
2003; Slabbekoorn & den Boer-Visser 2006). Other
traits might exhibit limited phenotypic plasticity, and
disturbances may result in population decline. Fur-
thermore, both domesticated and non-domesticated
animals residing in homes or zoos will demonstrate
behaviors that might no longer be adaptive. Research-
ers can design participatory projects in which volun-
teers collect data on species in different environments
and investigate different hypotheses behind complex
behaviors. In an online project called CamClickr, par-
209
ticipants added behavior-category tags to images
archived from nest cameras to help investigate varia-
tions in nest attendance (Voss & Cooper 2010).

Because funding cycles offered by granting institu-
tions tend to span three- to five-year periods, collect-
ing long-term data sets can be challenging for
research scientists. The collection of such data sets
could be made possible through citizen science activi-
ties. Depending on the initiative, initial investment to
set up the project, and maintenance costs, could be
low with free online tools for managing projects (e.g.,
www.citsci.org), recruiting volunteers (e.g., www.
scistarter.com), and data archiving (e.g., www.data-
one.org). Citizen scientists could measure different
components of fitness such as survival to maturity,
longevity, mating success, and reproductive success.
Phylogeny
Behaviors can also be understood from a phylogenetic
perspective. By considering ancestry, researchers can
identify traits that are highly conserved and those that
are more recently evolved within the same lineage.

Because closely related species are found in everyday
environments, a citizen science project could collect a
wide variety of behavioral data on closely related spe-
cies in different environments that have adopted dif-
ferent survival strategies. By comparing closely
related species occurring on different continents (e.g.,
black-billed bagpie Pica hudsonia in America and the
European magpie Pica pica in Eurasia), citizen scien-
tists can provide simultaneous data sets without the
confound of temporal displacement typical of smaller-
scale research projects.
Not all citizen engagement in animal behavior pro-
jects fits neatly into Tinbergen’s levels of analysis.
Recently, Project: Play With Your Dog (Horowitz Dog
Cognition Lab, Barnard College) requested short vid-
eos depicting play between a dog and a person, with
the aim of capturing the wide array of behaviors and
conditions that encompass interspecific play. Video

submissions drew from around the world, and
although participants did not contribute directly to
behavioral analysis, citizen scientists had to think
about behavior in terms of how they and their dog
play together and provide behavioral descriptors in an
accompanying questionnaire.
Similarly, an online survey by Animal Welfare Indi-
cators asked participants to assist in developing a con-
tinuous behavioral scale of lameness for goats.
Participants watched nine videos and scored abnormal
gait, head-nodding, and spine curvature on a visual
analogue scale. Before participants began, each
behavior had a clear description, and participants
were schooled in measuring behavior as per the speci-
fications of the project.
Merging Ethology and Citizen Science Methods
Creating Successful Citizen Science Animal Behavior
Projects

Like all successful research endeavors, the design of
citizen science projects follows the steps of the scien-
tific method; the only difference is the additional layer
of complexity introduced by the involvement of the
lay public. Projects start with the formulation of scien-
tific questions, organization of a team of skilled
experts in relevant areas, and determination of appro-
priate data collection (Bonney et al. 2009b). Depend-
ing on the legal status and various protections offered
to different species, researchers might need to present
protocols to Institutional Animal Care and Use
Committees (Bayne 1998). Citizen scientists can learn
behavior-coding techniques—or relevant project-
related skills—in online videos or in person. The
research team can control quality before the project
begins by testing each prospective participant’s
accuracy and ability to collect meaningful data.
Behavioral data accuracy is a key concern, and pro-
jects incorporating citizen science data should con-

sider topics like observation quality, sampling bias,
and interobserver reliability (Dickinson et al. 2010;
Burghardt et al. 2012; Hart et al. 2012). Williams
et al. (2012) found that although zoo visitor interest
in the project was high, the activity budget data col-
lected on a group of captive otters were not accurate,
mostly because visitors did not adhere to the observa-
tion period; the researchers suggest that certain
sampling methods might be easier—or harder—for
observers.
Meanwhile, there are numerous ways to improve
data quality in citizen science (Cohn 2008; Silvertown
2009; Wiggins et al. 2011; Bonter & Cooper 2012).
Technologies like smartphones, multimedia, or inter-
active video could assist in time-keeping and behav-
ioral data collection (Parr et al. 2004; Aanensen et al.
2009). Data management tools, often through cyber-
infrastructure, improve efficiency (Newman et al.

2011). Additionally, some citizen science projects
demonstrate that certain types of people collect more
accurate data than others (Delaney et al. 2008), and
similar lessons can make their way into citizen science
projects in animal behavior.
Finally, researchers can be creative in where and
how volunteers access animals and how behavior will
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be observed or documented. In some cases, behaviors
are not observed directly but are inferred. Phenology
research addresses the timing of life-cycle events,
including behaviors such as egg laying and migration
in birds, insects, and some reptiles and amphibians
(Head et al. 2013). Limited access to animals in situa-
tions where volunteers can obtain unbiased observa-
tions of life-cycle events may have been one of the

largest impediments to citizen science in animal
behavior. Now, Internet and remote-camera techno-
logies combined with crowdsourcing infrastructure
offer new opportunities to collect data on animals that
are difficult to access (Deng et al. 2012; Head et al.
2013; Loos & Ernst 2013). Live-streaming cameras
can be set up at areas with known activity—animal
nesting or gathering sites—so that people can docu-
ment behaviors at a distance. In addition to remote/
web-mediated studies, possibilities abound for studies
of companion animals (e.g., dogs, cats, rabbits, birds,
ferrets, fish, gerbils, and hamsters), zoo animals, ani-
mals viewed during nature tourism, and urban wild-
life such as squirrels, pigeons, rats, foxes, and coyotes.
Volunteers could document the behavior of animals
at zoos or wildlife tourism destinations, especially ani-
mals that handlers and keepers do not have time to
record. Special interest groups—such as the House
Rabbit Society, backyard chicken groups, and pet

breeders—offer significant potential for citizen science
projects.
Training and Resources for Animal Behavior Inquiry
Skill in observing and measuring animal behavior
does not necessarily follow from an appreciation of or
even cohabitation with animals. Members of the pub-
lic are generally interested in animals yet have a pro-
clivity not toward Tinbergen’s four levels of behavior,
but often toward anthropomorphic explanations of
behavior.
While the study of animal behavior is an attempt to
investigate ultimate and proximate questions about
behavior, humans often characterize animal behavior
through an anthropocentric lens (Foster et al. 2011).
People readily ascribe human characteristics to non-
humans. Subjective assessments of non-human
behavior can be consistent across observers, often
reifying existing beliefs regardless of whether they
have been substantiated (Bahlig-Pieren & Turner

1999; Morris et al. 2008; Horowitz 2011). Seminal
work by Heider & Simmel (1944) found that people
consistently attribute human emotions, aims, and
objectives to inanimate objects moving in a ‘human-
like’ manner. Companion animal species, mammals,
and non-human primates are more readily anthropo-
morphized than other species (Eddy et al. 1993), even
though seemingly ‘alien’ species, like cephalopods,
display complex behaviors and cognitive processes
(Fiorito & Scotto 1992; Moriyama & Gunji 1997).
Importantly, the appearance of behavioral homologies
should not imply similar meaning, such as the ‘smil-
ing’ chimpanzee or dolphin (Horowitz 2007).
Tendencies toward anthropomorphism are perhaps
a useful ‘hook’ when engaging the general public
in animal behavior projects. Recently, an online
video titled ‘Lol cats like stroking too’ (http://www.
youtube.com/watch?v=OFCRvjle2o8) accompanied a

publication on the neurons activated by massage-like
stroking to assist in public understanding of the
research (Vrontou et al. 2013). The video received
over 200,000 views in the first year.
Clearly defined project expectations and training
materials can minimize the effects of anthropomor-
phic inferences (Voss & Cooper 2010). Scientists can
provide a detailed ethogram, an explicit coding
scheme, and a clear behavior-sampling technique
(Martin & Bateson 2007). Behaviors in an ethogram
can be distinguished based on structural or empirical
characteristics, such as ‘raised lips at edges’, or conse-
quence-based descriptors, such as ‘threat’. Empirical
descriptors help create unambiguous behavioral
parameters (Lehner 1987) but might offer challenges
for unpracticed observers. Volunteers need a clear
understanding of the terms in which they are
expected to view and document behavior and have

the technology, such as a timer, to code appropriately.
Researchers determine who is being watched and
when and how to watch them. Some coding schemes
provide information on how much time an animal
spends performing various behaviors, while other
schemes monitor behavior frequencies (Altmann
1974). Researchers must match the skill level of
observers with the scientific question. The role of vol-
unteers in the study of animal behavior will largely
involve the production of meaningful behavioral data.
The challenge will be to offer effective guidance and
instruction to avoid anthropocentric assessments.
A number of institutions have created behavior-
observation instructional platforms. Ethosearch (www.
ethosearch.org), created by the Lincoln Park Zoo, is a
resource for researchers and students to store and view
ethograms and practice measuring animal behavior.
Living Links (www.living-links.org)—a joint enclosure

for squirrel monkeys (Saimiri sciureus) and capuchins
(Sapajus apella) at the Edinburgh Zoo—offers a
public-learning resource to observe and score the behav-
ior of both species (Macdonald & Whiten 2011). Cam-
211
Clickr was an online citizen science project to code the
behaviors of nesting birds from still images as a way of
carrying out scan sampling. Voss & Cooper (2010) cre-
ated a college-level laboratory unit in which students
can use CamClickr to create ethograms while learning
about observer bias, observation techniques, and defin-
ing and quantifying observations.
Conclusion
Animal behavior researchers might consider whether
and how citizen science could support and amplify
their studies. Volunteers can expand the scope and

scale of animal behavior studies by bringing attention
to novel or rare behaviors or providing access to
additional variation in animal behaviors. Volunteers,
too, can benefit from personal experiences engaging
in hands-on scientific interactions with animals.
Undoubtedly, challenges arise with the involve-
ment of citizen science in animal behavior, yet the
possibilities are great. Altmann’s (1974, p. 229) text
on behavior sampling methods stresses that there
should not be a disconnect between the laboratory
and the real world: ‘Such a restriction on research
would mean that the behavioral sciences would for-
ever forsake any hope of knowing whether their most
powerful theories have any relevance in the world of
behavior outside the laboratory.’ Citizen scientists
offer an additional, and untapped, opportunity for
ethological exploration.
Acknowledgements

Thanks to Felicity Muth, Mark Hauber, and Andrew
Fulmer for conceptual conversations. Many thanks to
reviewers for comments, which improved the manu-
script.
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