Propose one hypothesis that researchers studying animal behavior could examine using the help of citizen science volunteers.

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: julblue@gmail.com

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

ethologyinternational 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 GmbH208

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-

Ethology 120 (2014) 207–214 © 2013 Blackwell Verlag GmbH 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

Ethology 120 (2014) 207–214 © 2013 Blackwell Verlag GmbH210

Public Engagement in Ethology J. Hecht & C. B. Cooper

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-

Ethology 120 (2014) 207–214 © 2013 Blackwell Verlag GmbH 211

J. Hecht & C. B. Cooper Public Engagement in Ethology

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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