Introduction

Research in Zoos and Aquariums:

Purpose, Justification, Utility,
and Welfare
Michael Hutchins, Robert J. Wiese,
and Brandie Smith

The subject of zoo biology can also be briefly described as thus: it

embraces everything in the zoo which is biologically relevant.

Heini Hediger, Man and Animal in the Zoo (1969)

Zoological institutions are powerful wildlife conservation entities that

work to conserve wildlife and educate and inspire millions of people
about animals and their habitat each year (Gusset & Dick, 2010).
Professional accredited zoos and aquariums are required to actively
engage in conservation and be committed to research and scientific
advancement (AZA, 2018; Barongi, Fisken, Parker, & Gusset, 2015).
These activities have increased exponentially in the past few decades,
and conservation, research and education rival recreation and visitor
engagement in the list of zoo and aquarium priorities (Conde, Flesness,
Colchero, Jones, & Scheuerlein, 2011; Conway, 1969, 2003; Field &
Dickie, 2007; Hutchins & Conway, 1995; Hutchins & Smith, 2003;
Konstant, 1995; Mallinson, 2003; Maple, 2016; Rabb & Saunders, 2005;
Zimmerman, 2010; Zimmerman & Wilkinson, 2007).
Over the past few decades, many authors have discussed various
aspects of zoo- and aquarium-based research (e.g., Anderson, Maple, &
Bloomsmith, 2010; Beck, 1974; Benirshke, 1975, 1996; Chiszar, Murphy,
& Smith, 1993; Hediger, 1964, 1968, 1969; Hofer, 2011; Hutchins, 2010;
Hutchins, Dresser, & Wemmer, 1995; Jarvis, 1967; Kaufman &
Zaremba, 1995; Lawson, Ogden, & Snyder, 2008; Lindburg, 2008;

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Marlieve & Newman, 1995; Maple, 1982, 2008, 2016; McCormick-Ray,

1993; Schwartz, 2017; Wharton, 2007, 2008). The goal of this chapter is
to provide a broad overview of key issues related to the history, purpose,
justification, utility, and diversity of research conducted in modern
zoological parks and aquariums (hereafter “zoos”) today. We will also
touch on a variety of important issues associated with the conduct of
research in zoological institutions, including a brief exploration of the
history of zoo research; advantages and disadvantages of conducting
research in the zoo setting; the diversity, purpose, and utility of zoo
research; administration of zoo research programs; research priority
setting; zoo–university and other partnerships; animal welfare and other
ethical considerations; publication of research results; and funding to
sustain zoo research programs.

   

Wemmer and Thompson (1995) and Wharton (2007) both provide
summaries of the history of zoo research worldwide. As the authors
point out, most scientists and many zoo staff have not viewed zoos as
research institutions, but a surprising amount of research is con-
ducted in zoos and published in mainstream scientific journals and
books today (see “Diversity of Research Topics” section). The pres-
ence of live animals, many of them little known to science or difficult
to observe in nature, and of people seeking to gain new knowledge or
experiences makes zoos a rich laboratory for scientific study.
Heini Hediger’s (1969) book Man and Animal in the Zoo was a
milestone in the history of zoo-based research. It was here that the
author coined the term “zoo biology,” referring to the study of wildlife
and humans in the zoo setting. However, research played a prominent
role in some early zoos. For example, the charter of the Zoological
Society of London, established in 1826, aspired to assemble a collec-
tion of living animals that could be used for scientific research instead
of just as a visitor attraction (Olney, 1980). The New York Zoological
Society was established in 1895, and by 1903, William Beebe, a young
curator at the Bronx Zoo, was already engaged in field expeditions to

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study wildlife in nature. By the mid-twentieth century, some zoos had

officially recognized research as part of their core mission. Spurred on
by the extinction crisis in the 1960s and 1970s (the US Endangered
Species Act was signed into law in 1972), by the 1980s and 1990s,
many zoos included research as part of their mission statements and
organized programs were beginning to develop, with formal research
protocols and dedicated personnel.

  

  
There are many advantages and disadvantages to conducting research
in zoos (Hutchins, 2001). Animal protectionists (e.g., Jamieson, 1985)
are often quick to point out the disadvantages, which can be substan-
tial when animals are maintained in inadequate social or physical
environments. For example, development, behavior, physiology, and
reproduction can all be modified – sometimes severely – under the
conditions of captivity, especially when the environment, including
size and quality of space, group composition, and dynamics and diet,
does not meet an animal’s basic biological or behavioral needs
(Hediger, 1964, 1968, 1969; McPhee & Carlstead, 2010; Meyer-
Holzapfel, 1968; Morris, 1964). This, in turn, can call into question
the validity of research results (Hosey, 1997; Hutchins, 2001). In the
past, inadequate environments have resulted in a wide range of prob-
lems, including, but not limited to, stereotypic behavior, inactivity,
and poor physical condition, often leading to stress, disease, shortened
life spans, and reproductive failure (Hediger, 1964, 1968, 1969; Meyer-
Holzapfel, 1968; Morris, 1964; O’Regan & Kitchener, 2005).
Fortunately, as zoos’ knowledge of wildlife biology, ecology, and
animal care and welfare has increased, enclosures have become larger
and more naturalistic and complex (Coe & Dykstra, 2010; Hutchins,
2003; Maple, McManamon, & Stevens, 1995), and as animal managers
and scientists have paid increasing attention to species-appropriate
environmental enrichment (e.g., mammals: Hoy et al., 2010), includ-
ing the composition of social groups and feeding strategies and

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nutrition, conditions for many zoo animals have greatly improved

(e.g., great apes: Hutchins et al., 1991), thus making the zoo setting
even more appropriate for scientific study (Hosey, 1997; Wemmer &
Thompson, 1995).
Jamieson (1985) questioned the appropriateness and utility of
zoo-based research. His statements that “research that is conducted
in zoos can be divided into two categories: studies in behavior and
studies in anatomy and physiology” (p. 112) and that the latter are
“the most common forms of zoo research” have, in recent decades,
been completely disproven, and, in retrospect, were not even accurate
at the time of publication. Based on this, he further concluded that the
benefits of zoo-based research do not outweigh the “moral presumption
against keeping animals in captivity” (Jamieson, 1995, p. 53). However,
Hutchins, Smith, and Allard (2003) countered his and Regan’s (1995)
arguments, demonstrating how zoo and aquarium commitments to
their primary missions of wildlife conservation and animal care/welfare
can overcome this presumption and “provide a powerful ethical justifi-
cation for accredited zoos and aquariums” (p. 964).
The advantages of zoo research are many. First, access to a wide
variety of species, many of which are little known to science, can be a
tremendous advantage for researchers seeking to contribute to our
knowledge. For example, little is known about many aspects of the
basic biology of arboreal, nocturnal, fossorial, aquatic, mountain-
dwelling, or other species that are difficult or impossible to study in
nature, but can be studied in the zoo setting (Hutchins, Dresser, &
Wemmer, 1995). In such cases, virtually any data collected can be new
to science and potentially applied to their captive management or
conservation in nature.
Second, from a practical viewpoint, zoo animals are found in
roughly the same location daily and are thus consistently available for
study. They are also comparatively easily observed, although the
advent of larger, naturalistic enclosures has, in some cases, made this
more difficult. This is also true of radio-tagged animals in nature, but
because they can move over long distances, they may still be difficult

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to find or otherwise hard to observe during certain phases of their life

cycles. For example, Hutchins (1984) studied an introduced popula-
tion of free-ranging Rocky Mountain goats (Oreamnos americanus) for
four years in Olympic National Park, Washington, but was never able
to observe a birth. The species inhabits rugged, high-altitude terrain,
and females typically isolate themselves during birthing, making
observations difficult. However, he was able to observe and document
several births at the Woodland Park Zoo in Seattle and to record early
postpartum behavioral development, thus contributing to our know-
ledge of this species (Hutchins et al., 1987).
Animals can also be easily found and observed in laboratory
settings, but in these cases, the environments – typically small, sterile
cages, which may facilitate large sample sizes and experimental and
control populations – are not ideal for animals, meeting only their
basic biological needs. While these conditions may be appropriate for
physiological studies or to experimentally test reactions to medicinal
drugs, they would be, in most cases, of only limited utility to meet the
goals of most contemporary zoo research. The fact that modern zoos
seek to keep animals in larger, more appropriate enclosures and social
groups essentially makes them intermediate between laboratory and
field settings.
In zoos, animals can be more easily captured or trained in order
to collect biological samples, such as feces, urine, blood, milk, glan-
dular secretions, and hair, or to take physiometric measurements
documenting growth and development though ultrasound or other
methods compared with in nature (Mellen & McPhee, 2010). Captures
of free-ranging wildlife, even through the use of immobilizing drugs,
can be stressful and result in injury or death (e.g., La Grange, 2006).
Sample collection in zoos is often done during routine health exams or
from closely monitoring animals and then collecting samples nonin-
vasively from known individuals. In some cases, such as blood or
urine sample collection, animals can be trained to present themselves
for the procedure, a non-stressful method for which the subject is
positively rewarded (Mellen & McPhee, 2010). In some cases, using

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these procedures themselves presents many opportunities for both

research (Schuele, 2016) and education (Lukas et al., 1998).
The age, sex, and genealogy of individual zoo animals and their
reproductive histories are often known, which is a tremendous advan-
tage for certain kinds of research (Schwartz, 2017). Under field condi-
tions, this information is often unknown or difficult to come by
unless animals can be marked or otherwise identified for long periods
of time.
There are disadvantages to zoo-based research as well. Although
one goal of modern zoo animal management is to give animals oppor-
tunities to express as large a portion of their natural behavioral reper-
toires as possible (Hutchins, 2003; McPhee & Carlstead, 2010), captive
animals are kept in environments that may resemble, but do not repli-
cate exactly the natural habitats the species frequents in nature. It is
important to note, however, that conditions in nature can vary greatly
as well (e.g., elephants: Hutchins, 2006a), making it difficult to account
for such variability when designing zoo management programs.
In many cases, research opportunities are limited by practical
animal management or exhibition needs (Hutchins, 1988; Kleiman,
1996). For example, conspicuous marking to identify individual
animals may be prohibited by zoo management, as it detracts from
the visitor experience. Furthermore, in order to prevent unwanted
reproduction, males are sometimes isolated from females when the
latter are in reproductive condition, thus limiting some research oppor-
tunities. In addition, not surprisingly, predators and prey are not kept
together in the same enclosures, thus limiting the expression of preda-
tor and anti-predator behavior. Similarly, group composition or history
may not be what it is in nature, as, for example, in African elephants,
where herds normally consist of several generations of related females
(Poole & Moss, 2008). Although this is changing (Hutchins,Smith, &
Keele, 2008), given limited availability, zoos typically have had to form
elephant groups from unrelated animals. This could alter social behav-
ior, but elephants have also shown the ability to integrate into these
groups and form attachments to unrelated individuals.

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One obvious disadvantage of zoo-based research is small

sample size, which can make it difficult to draw statistically sig-
nificant conclusions (Kuhar, 2006; Plowman, 2008). This is espe-
cially true of rare or endangered animals that are seldom exhibited
in zoos or may be present in limited numbers. That being said,
this disadvantage can sometimes be overcome through coopera-
tive studies that extend over several institutions, thus greatly
expanding the number of subjects involved (Kleiman, 1996). For
example, a study investigating the impact of dominance status on
ovarian activity in female African elephants utilized 33 subjects
housed at 14 different facilities across North America (Freeman,
Schulte, & Brown, 2010).
It should also be noted, however, that when little or nothing is
known about a species, even studies based on small sample sizes can
make important contributions. Consider, for example, Ogden, Olson,
and Miner, (1991) study of a single pair of golden monkeys at Seattle’s
Woodland Park Zoo and Portland’s Washington Park Zoo. Another
example is Plair, Reinhart, and Roth, (2012) study of neonatal behav-
ioral milestones and growth in two Sumatran rhino calves born at the
Cincinnati Zoo. Virtually nothing was known about these species’
reproductive biology, behavior, or development at the time, so these
studies made contributions of scientific significance.
Another possible disadvantage is the lack of trained personnel
(Anderson et al., 2010). In order to conduct and publish valid scientific
research, zoos must have trained scientists on staff or otherwise
develop cooperative relationships with academic institutions or non-
governmental organization partners that possess this expertise
(Fernandez & Timberlake, 2008; Hutchins, 1988; Kleiman, 1996). This
is not to say that zoo staff, including keepers and curators, and volun-
teers cannot be trained to contribute to scientific research – they
certainly can (Hutchins, 1988; Kleiman, 1980; Whitham & Wieleb-
nowski, 2009) – but supervision by trained scientists with necessary
skills, such as study design, statistics, interobserver reliability, and
publication, is critical for long-term success.

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Unlike university research laboratories, where controlled experi-

mental studies are possible, zoos seldom conduct manipulative experi-
ments on their animals, preferring instead to rely on naturalistic
observations (which may include quantitative methods) to collect
data. While this may be a disadvantage for determining causal rela-
tionships between variables, it is still contributing to our overall
knowledge of a species’ basic biology and behavior. In addition, it is
far superior to and more efficient than relying on simple trial and
error, which is what zoos did in the past (Hutchins, 2001). It should
be noted that field studies seldom provide opportunities for manipu-
lative experimental studies either. That being said, some manipula-
tions may be possible, which provide the conditions of natural
“experiments.” This can involve before-and-after studies of subjects
under different conditions. For example, Meller, Croney, and Shep-
herdson (2007) studied the effects of rubberized flooring on elephant
behavior at the Oregon Zoo. The authors took advantage of changes
that were occurring in indoor elephant holding at the institution, first
collecting baseline data when the animals were on traditional con-
crete floors and then when the rubberized floors were installed. They
found increases in locomotion and normal sleep behavior (standing vs.
recumbent) and decreased discomfort behaviors on the rubberized
floor when compared with the concrete floor, suggesting the new
flooring was a welcome addition. Studies on diet are also a common
way to get before-and-after comparisons in the zoo environment (see
Chapter 10, this volume).

     :

  
Several authors have attempted to summarize the scope of research
in modern zoos and aquariums (Anderson, Keiling, & Maple, 2008;
Finlay, James, & Maple, 1986; Hardy, 1996a, 1996b; Hutchins, Paul, &
Bowdoin, 1996; Kaufman & Zaremba, 1995; Kleiman, 1992; Maple
& Bashaw, 2010; Maple & Finlay, 1989; Melfi, 2007; Stoinski, Lukas,
& Maple, 1998; Wemmer, Rodden, & Pickett, 1997). The diversity of

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topics studied in the modern zoo is mind-boggling. In this section, we

review some of those topics and provide a few examples and justifica-
tions based on their utility. This is in no way a comprehensive over-
view, as we have left out numerous topics in the interest of brevity.
Research is often categorized according to its goals. “Applied”
research is focused on problem-solving. For example, any research
aimed at improving zoo animal management or husbandry would be
classified as applied research. In contrast, “basic” or “pure” research
is intended to improve our understanding of the natural world,
irrespective of its possible practical application (Hutchins, 2001;
Thompson, 1993).
Most zoo-based research is applied. However, basic research is
conducted as well. For example, elucidating why Caribbean flamingos
(Phoenicopterus ruber) tend to stand on one leg (Anderson &
Williams, 2010) is a basic research question, with little or no immedi-
ate practical application to animal management or conservation,
but that could have theoretical or scientific significance. Another
example was the discovery by the National Aquarium in Baltimore
and Baltimore Zoo that toxic alkaloids in the skin of poison dart
frogs (Dendrobatidae) are not present in captive-bred individuals and
have a dietary origin (Daly et al., 1994). Still another would be hand-
edness (laterality) in primates (e.g., ring-tailed lemurs: Hosey, Hill, &
Lherbier, 2012). That being said, it is important to realize that pure
research conducted today may have practical value in the future, even
though its usefulness may not be immediately evident (Hutchins,
2001; Hutchins & Thompson, 2008; Thompson, 1993).

Anatomy and Physiology

Zoos can be valuable collaborators to assist with studies of the basic
anatomy and physiology of animals little known to science. Living
animals make it possible to study a wide variety of topics, includ-
ing blood chemistry and characteristics, which are essential for estab-
lishing “normal” health parameters. This, in turn, can help veterinarians
determine when an animal is ill or in poor condition. For example,

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Ange-van Heugten, Verstegen, Ferket, Stoskopf, and van Heugten, (2008)

studied blood serum chemistry concentrations of 30 captive wooly
monkeys (Lagothrix lagotricha) housed at two European institutions as
a means of establishing normal parameters to monitor health. Common
causes of death in this species are pregnancy and hypertension, thus
making these data useful for health evaluation. Similarly, Abbondanza,
Power, Dickson, Brown, and Oftedal, (2013) were able to investigate
variation in milk composition of three lactating female Asian elephants
(Elephas maximus) from birth to the calf’s third year of age and to
determine changes that occur over time. Such studies would be difficult,
if not impossible to conduct on free-ranging elephants.
All living animals will someday also die, and animals are often
valuable from a scientific perspective even after their deaths (Hutch-
ins, 2003). While anatomical studies are not typically conducted by
zoos themselves, zoos are often a source for animal carcasses/skel-
etons for museum study collections or university anatomists who
wish to better understand the anatomical structures of various species
that may be little known to science (Hutchins, 1990).

Animal Welfare
The issue of how to assess animal welfare – or the well-being of
individual animals – housed in zoos is a hot scientific topic and one
that is in its early stages of development. There are, in fact, many
different philosophies and scientific approaches to the study of animal
welfare, recently summarized by Barber (2009), Fraser (2009), Hill and
Broom (2009), and Veasey (2017). This will undoubtedly be an import-
ant and controversial topic of investigation for the coming decades.

Behavior
Behavioral research is a common focus of zoo research and many
authors have discussed its role and utility (Eisenberg & Kleiman,
1977; Forthman & Ogden, 1992; Hediger, 1968; Hosey, 1997; Kleiman,
1992; Lindburg, 2010; Melfi, 2005; Moran & Sorensen, 1984). Behav-
ioral research not only improves our understanding of a species’ basic
behavioral biology, but also helps to improve care and husbandry

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(e.g., Eisenberg & Kleiman, 1977; Kleiman, 1994; Lindburg, 2010;

Lindburg & Fittch-Snyder, 1994). Developing successful manage-
ment strategies in the absence of such information can be difficult,
if not impossible.
Considerable research has, for example, been conducted on the
parental care behavior of endangered golden lion tamarins (Tardie et al.,
2002), contributing greatly to their management in zoos. Males and older
sibling “helpers” have proven to be essential in the care of infants, and
monogamy is the prevalent mating strategy. This has had huge implica-
tions for the composition of managed groups as it relates to reproductive
success. Similarly, research at the Bronx Zoo found that mate choice was
a very important aspect of successful mating and reproduction in lekking
bird species, such as the red bird of paradise (Hundgen, Hutchins, Shep-
pard, Bruning, & Worth, 1991). In this case, reproduction was most
successful when females were given an opportunity to choose between
alternative mates, as they would in nature. Giant panda husbandry and
management, too, have benefited greatly from behavioral studies (Zhang,
Swaisgood, & Zhang, 2004), as have a wide variety of taxa.

Cognition
One aspect of behavioral research that is important from both an
applied and theoretical perspective is animal cognition. What kinds
of cognitive abilities do non-human animals have and how does these
adapt them to their specific lifestyles and ecological niches? Cognitive
abilities also have implications for captive management, in that
species with high-level cognitive function often require management
strategies that reduce boredom and enhance complexity and control
(Clark, 2016; Clark, F. E., Davies, S. L., Madigan, A. W.,Warner, A. J.,
& Kuczaj II, 2013). Zoos are excellent places to test the cognitive
abilities of animals, such as elephants (e.g., Plotnik, de Waal, Moore,
& Reiss, 2010), great apes (Krupenye, Kano, Hirata, Call, & Toma-
sello, 2016), and dolphins (see also Chapter 19, this volume). Such
studies, often combined with field observations, have resulted in
significant contributions to our understanding of animal cognition
(e.g., dolphins: Pack, 2010; Wells, 2009).

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Contraception
The practice of contraception (the prevention of pregnancy, usually
though artificial means) has become increasingly important to zoos
that seek to manage their populations within available space while
also maintaining genetic diversity (Asa & Porton, 2005, 2010; Hutch-
ins, 2003). Breeding animals randomly and without control is detri-
mental to both genetic and demographic management (Ballou et al.,
2010). Furthermore, the production of surplus animals creates many
ethical dilemmas (Asa, 2016; Lindburg & Lindburg, 1995). Thus, it
is no surprise that zoos have invested considerable resources into
research on the development of safe and effective hormonal contra-
ception methods; the results of much of this research were summar-
ized in Asa and Porton (2005, 2010).

Disease, Pathology, Veterinary Care, and Mortality

Conservation medicine will be a major emerging area of priority
research in the coming years (Vitali, Reiss, & Eden, 2011) as climate
change exacerbates the spread of mosquitoes and other disease vectors
(Barbosa, 2009; Macek, 2014). For example, avian malaria and West
Nile virus are two known pathogens that can and do impact both free-
ranging and captive bird populations (Nemeth & Ossterle, 2014). The
latter was discovered in North America by a pathologist at the Wild-
life Conservation Society’s Bronx Zoo in New York City (Lanicotti
et al., 1999).
Studies of diseases and parasites afflicting zoo animals are also
important for monitoring health and treating various medical prob-
lems. For example, Poynton and Whittaker (1994) documented the
protozoa present in the intestines of wild-caught and captive-bred
poison dart frogs (Dendrobatidae) from the National Aquarium in
Baltimore. The authors’ findings supported the judicial use of anti-
protozoal agents to keep these organisms from becoming pathogenic.
Retrospective studies focusing on the reasons for species-specific
mortality can have important implications for captive management

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and husbandry. For example, Mendyk, Newton, and Baumer, (2012)

investigated the reasons for mortality in varanid lizards at the Bronx
Zoo in New York. The authors found bacterial infections, neoplasia,
female reproductive disorders, gout, and endoparasitism to be the pri-
mary causes of death and recommended several improvements in man-
agement to enhance survival.

Education/Visitor Studies
Zoos have touted themselves as educational facilities, the goal being
to teach the public about animals and their conservation (Anderson,
2003; Ogden & Heimlich, 2009; Rabb & Saunders, 2005; Sterling,
Lee, & Wood, 2007; Thomas, 2015). But how effective are they at
disseminating such information and having the public retain it or
take action? These are legitimate research questions that beg to be
answered (Moss, 2015; Moss & Essen, 2013). Critics have questioned
the educational role of zoos, so it is important that zoos be able to
counter with evidence from legitimate scientific studies. While
some studies have indicated questionable impacts or even a lack of
effectiveness (Balmford et al., 2011), others have documented both
learning and retention (Falk et al., 2007; Jensen, Moss, & Gusset,
2017). Still others have looked at specific visitor reactions to animals
or exhibits. For example, Miller (2012) studied visitor reactions to
films showing captive animals pacing and found that it had a detri-
mental effect on their perceptions of the quality of animal care.
Others have looked at how zoo visitors reacted to various exhibits
at Chicago’s Brookfield Zoo (Luebke & Matiasek, 2013) or investi-
gated factors that influenced zoo visitors’ attitudes toward wildlife
and nature conservation (Swanagan, 2000).

Environmental Enrichment
Shepherdson (1999) defined environmental enrichment as “an
animal husbandry principle that seeks to enhance the quality of
captive animal care by identifying and providing the environmental
stimuli necessary for optimal psychological and physiological

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well-being” (p. 1). While environmental enrichment takes many

forms – both naturalistic and artificial – its effectiveness at produ-
cing the desired results is a legitimate and important topic of
research for modern zoos and laboratories that house and care for
living animals. Much of the existing research at the time was sum-
marized in Shepherdson, Mellen, and Hutchins (1999), but relevant
studies have proliferated in recent years. For example, Clark and
Melfi (2012) studied the effect of feeding and sensory environment
enrichment on the behavior of nine-banded armadillos, Senegal
bushbabies, and two-toed sloths housed in a multi-species display
in the nocturnal house at Paignton Zoo in the UK. They found that
feeding and scented pine cones had a positive effect on the animals,
increasing behavioral diversity.

Enclosure Utilization and Activity Patterns

Knowing how animals utilize their exhibit space and when and
where they are most and least active is important for providing
quality care. For example, Blowers, Waterman, Kuhar, and Bettinger
(2012) investigated enclosure utilization of a group of nine female
Nile hippopotamus (Hippopotamus amphibious) in a large, natural-
istic exhibit at Disney’s Animal Kingdom. Hippos aggregated in
preferred areas of the exhibit, mostly in water at depths of 0.6–1.0
meters. The authors suggested that their findings could be useful for
naturalistic exhibit design for this species.
Activity patterns can also be an indicator of the appropriateness
of the zoo environment and may indicate the need for modification.
Challender, Thai, Jones, and May (2012) studied time budgets and
activity patterns in Suna pangolins (Manis javanica) housed at the
Carnivore and Pangolin Conservation Programme Facility in Viet-
nam. They found individual variation in how animals partitioned
their active time, but that most activity occurred between 18:00 and
21:00 hours. Two animals exhibited excessive pacing and clawing
activity, thought to be stress related; cessation of this behavior in
one individual occurred with enclosure modification.

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Field Research
As zoos have gotten increasingly involved in field conservation
activities (Conway et al., 2003; Hutchins & Conway, 1995; Zimmer-
man, 2010; Zimmerman & Wilkinson, 2007), they have also become
involved in associated field research on wildlife and their habitats.
Organizations such as the zoo-based Wildlife Conservation Society,
London Zoological Society, Durrell Wildlife Conservation Trust, and
Frankfurt Zoological Society have long histories of support for in situ
wildlife research and conservation (Wemmer & Thompson, 1995;
Wharton, 2008). Others have come on board more recently. Many
examples of field research by zoos are summarized in Conway et al.
(2003), including studies of the biology and ecology of the Aruba
Island rattlesnake (Odum, 2001; see also Odum & Reinert, 2015)
and the Jamaican iguana (Hudson, 2001), as well as rehabilitation
of spawning habitat for intertidal fish in British Columbia (Marlieve
& Matrtell, 2001), just to name a few.

Genetics
Knowing the genetic relationships of animals in small, managed
populations is key for maintaining genetic diversity over time (Bal-
lou et al., 2010). Normally, genetic relationships are inferred by
animal records documenting parental and other familial relation-
ships. However, human error can occur in record-keeping, and
research identifying parentage through genetic markers can be
important for improving accuracy. Ferrie et al. (2013), for example,
used molecular markers to confirm parentage in a captive flock of
Marabou storks (Leptoptilos crumeniferus), thus improving the
accuracy of studbook records. Genetic studies can also be used to
determine gender in taxa that lack external genitalia or other sec-
ondary sexual characteristics, like some birds. For example, Vucice-
vic et al. (2013) evaluated the CHD gene as a universal molecular
marker to sex birds in 58 avian species; gender identification was
successful in 50 of those species.

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Emerging genetic technologies such as stem cell technology and

genome editing are being investigated to bring back lost genetic vari-
ation from stored tissue banks and to add disease resistance to suscep-
tible populations. San Diego scientists have also produced pluripotent
stem cells from frozen tissue in the beginning steps to create embryos
of the northern white rhino that is functionally extinct in the wild
(Saragusty et al., 2016).

Growth and Development

Studies of growth and development require regular observations or
measurements to document changes that occur over time. Such
studies are much easier when identifiable animals can be observed
or handled on a regular and predictable basis from birth to adulthood.
Zoos are, in fact, the ideal venue in which to conduct growth and
development studies, as such long-term regularity of sampling is
often difficult, if not impossible in the field. As has been mentioned
for other kinds of research, however, a combination of information
coming from field and captive studies provides a more complete
picture of this aspect of a species’ behavior and biology than one
information source alone.
One such study utilized a survey to assess the physical and
behavioral development of 12 African elephant calves from birth to
3 months of age across 7 different facilities in North America and
Europe (Kowalski, Dale, & Mazur, 2010). Another examined devel-
opmental milestones among African elephant calves on the first day
of their lives, combining data from both field and captive situations
(Bercovitch & Andrews, 2010). The presence of advanced medical
equipment in zoos, such as ultrasound, has opened up incredible
research opportunities for in-womb developmental studies. Drews
et al. (2011), for example, were able to use sonography to record the
development of seven bonobo (Pan paniscus) fetuses from four preg-
nant females during gestation at the Milwaukee County Zoo. The
females were trained to present themselves for examination using
positive reinforcement techniques.

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Husbandry and Care

Much zoo research has been focused on improving animal care and
husbandry (e.g., Eisenberg & Kleiman, 1977; Hutchins et al., 1995;
Lawson et al., 2008; Melfi, 2005), and this can involve a better under-
standing of many of the topics covered here. This is appropriate as
zoos wish to constantly improve the ways they care for animals and to
sustain their populations for both exhibition and other purposes,
including research, education, and conservation (Hutchins, 2003;
Hutchins & Smith, 2003). This can be a huge challenge when little
or nothing is known about a species’ basic biology or behavior in
nature. That being said, best practices for zoo managers often arise
from a combination of knowledge gained from captive and field stud-
ies, which leads to a more complete picture of a species’ basic biology
and behavior (e.g., dolphins: Dudzinski, 2010; Samuels, 1996).
Some of the best known examples of zoo-based research contrib-
uting to animal care and husbandry have been conducted on large
mammals, such as giant pandas, rhinoceros, and tigers. The breeding
and care of these taxa have been greatly improved through both zoo-
based and field research, which has led to a more detailed understand-
ing of their basic behavior, biology, and management, as summarized
by Lindburg and Baragona (2004), Hutchins and Kreger (2006), and
Tilson and Nyhus (2010), respectively. However, many other taxa
have benefited as well, including golden lion tamarins (Kleiman &
Rylands, 2002), Komodo dragons (Murphy, Ciofi, de la Panouse, &
Walsh, 2002), cranes (Ellis, Gee, & Mirande, 1996), and black-footed
ferrets (Miller, Reading, & Forrest, 1996), just to name a few.

Longevity
How long individual animals live is an interesting research question.
Longevity is theoretically interesting in that senescence is an evolved
trait. However, the lives of individuals can be greatly modified by a
variety of different factors, such as condition, reproductive effort,
disease, and so forth. From a practical perspective, critics have often

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used maximum longevity as a reason for attacking zoos, claiming that

zoo settings result in early death when compared with free-ranging
animals (see Borrell, 2008). However, this is misleading, as population
data measure average life spans and standard deviations indicate vari-
ation. Countering such misinformation through science can therefore
be useful. For example, Robeck, Willis, Scarpuzzi, and O’Brien (2015,
p. 1) compared life history parameters in killer whales (Orcinus orca)
and concluded that the study provided “evidence for similar life-
history parameters of free-ranging and captive killer whale popula-
tions and the reproductive potential and survivorship patterns estab-
lished herein have application for use in future research concerning
the overall health of both populations.” Wiese and Willis (2004) stud-
ied longevity in elephants and found similar patterns in captive versus
free-ranging elephant populations.

Media Studies
Studies of the media’s treatment of zoos – whether deserved or not –
can be important for formulating strategies for dealing with reporters.
Crisis management is certainly an issue that zoo mangers must deal
with frequently during their tenures. Science can inform this process.
For example, Hutchins (2006b) studied the way that the media
responded to zoo animal deaths. From a sample of stories collected
randomly from the Internet, he was able to categorize and quantify
the ways that the media responded to zoo animal deaths, ranging from
accusatory to neutral to sympathetic. The results had important
implications for the way that zoo public relations staff characterized
zoo animal deaths with the media. He concluded, “A greater focus on
the science of zoo animal death is necessary for accredited zoos to
maintain the public’s confidence in their animal care practices.” (p. 1).

Nutrition and Diet

Determining a viable captive diet for exotic species can be difficult, as
food items utilized in nature are not typically available at the local
grocery store (Crissy, 2005). Much research has therefore examined

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issues such as the nutritional and energy content or digestibility of

artificial diets, which can help to determine the amount, type, and
appropriateness of food offered. For example, Clauss, Kleffner, and
Kienzle, (2010) studied the digestibility and nutrient content of diets
fed to a wide variety of carnivores. Similarly, Bennett, Booth-Binczik,
and Steele, (2010) investigated the nutritional content and digestibility
of a commercially available diet fed to ocelots (Leopardus pardalis).
Dietary studies are particularly important for animals that have
specialized diets in nature, such as obligate insectivores. For example,
Finke (2015) found that commercially available insects used in zoo diets
lacked several key nutrients and thus needed vitamin supplementation
through dusting or gut-loading (e.g., Michaels, Antwis, & Preziosi,
2015). Clark, Silva-Fletcher, Fox, Kreuzer, and Clauss (2016) surveyed
UK zoos to study feeding practices, body condition, and feces consist-
ency in captive insectivores, such as giant anteaters. How and how often
food is presented are other important research questions that can have
important implications for stimulating activity in many species in cap-
tivity (Lindburg, 1998) (see “Environmental Enrichment” section).

Reintroduction
Zoos sometimes breed animals for reintroduction into the wild to aid
their conservation in nature (Beck, Rapaport, Stanley Price, & Wilson,
1994; Earnhardt, 2010; Gilbert & Soore, 2017; Price & Soore, 2003;
Stanley-Price & Fa, 2007). While reintroduction of captive-bred wildlife
is not a common occurrence and has its challenges, it has helped
species as diverse as the American and European bison, Arabian oryx,
black-footed ferret, Pere David’s deer, California condor, Kihansi spray
frog, whooping crane, and northern bald ibis, just to name a few (see
Chapters 13 and 15, this volume, for the stories of some of these).
In such cases, research has been critical for determining rearing,
pre-release conditioning, and release strategies for animals targeted
for reintroduction (e.g., hard vs. soft releases). For example, research
on pre-release conditioning for black-footed ferrets at the Smithsonian
Institution’s Conservation Biology Institute in Front Royal, VA, led to

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the development of successful training techniques to allow naïve,

captive-bred ferrets to recognize and kill their primary prey of prairie
dogs, escape danger in burrows, and recognize predators, experiences
that helped released animals survive in nature (Miller et al., 1998).

Reproductive Biology
Research into animal reproductive biology is a common occurrence in
zoos due to its relationship with successful breeding and reproduction
(Asa, 2010; Spindler & Wildt, 2010). In particular, an understanding
of a mammalian species’ basic reproductive physiology, including
female estrus cycles, is critical for success. A classic example is
SeaWorld’s work on the female hormonal cycle of killer whales
(Orcinus orca), which was determined through analysis of urine
samples collected from captive animals trained to urinate on cue
(Robek et al., 1993). As it involved regular sample collection, these
are data that would have been next to impossible to obtain from free-
ranging individuals and have contributed greatly to our understanding
of the species’ basic reproductive biology.

  

In order to be useful (i.e., to solve particular problems in animal care,
husbandry, or conservation – applied research) or to contribute to our
basic knowledge of animals or their habitats (basic or pure research),
research must be strategic (Hutchins & Thompson, 2008). This means
careful planning and consideration of what to study and why.
Developing a list of research priorities could come from exam-
ining the literature to see what has been done previously or by speak-
ing to animal managers (e.g., curators, keepers) to understand what
kinds of problems exist and therefore what kinds of research would be
useful to them. These types of gap analyses help us understand the
nature and variety of research needs that exist in any chosen field of
study. The expertise and interest of the researcher are also relevant, as
research projects take a great deal of time and effort to complete and
scientists should be motivated to take their project to completion.

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An example of just such a gap analysis comes from Melfi (2009)

for environmental enrichment research in zoos. The author surveyed
the literature and found that most enrichment studies to date
were focused on large mammals, notably primates, bears, large
cats, and elephants. She further pointed out the need to work with
a wider variety of taxa, something that was also stressed by Mellen,
Shepherdson, and Hutchins (1999).

Zoo Research Program Administration

Zoos that engage in research activities – either by in-house staff or by
visiting scientists – need to be organized from an administrative
perspective (Eisenberg, 1975; Hutchins, 1988; Kleiman, 1992, 1996).
Zoos must determine what kinds of studies are appropriate for
the zoo setting, develop criteria for evaluating research proposals
(Hutchins, 1988; Kleiman, 1985; Schaeffer, 1996), and also have
access to the results when studies are completed. This means having
a research policy in place, as well as some sort of program and
personnel to monitor projects, track and store the results, ensure
that the results are seen and perhaps utilized by zoo staff, approve
publication and public relations decisions, administer outside bio-
logical materials requests (Hutchins, 1990), and manage an animal
welfare committee to weigh the advantages and disadvantages of
research that might harm animals (see “Welfare and Other Ethical
Considerations” section). The absence of policy and procedure could
result in chaos, in safety concerns (especially when outside research-
ers are allowed into off-exhibit areas), in the loss of information, or
in public relations disasters, all issues that public or private insti-
tutions would like to avoid whenever possible (Hutchins & Ballen-
tine, 2001).

Zoo–University Relationships
One way zoos can develop a research program at little cost or to
augment their existing internal research efforts is to develop a
formal, cooperative relationship with a local college or university

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(Anderson et al., 2010; Eisenberg, 1975; Feistner & Price, 2002;

Fernandez & Timberlake, 2008; Hutchins, 1988; Lawson et al.,
2008; Lukas & Maple, 1997; Savage, 1988). Colleges and universities
have trained scientists on staff as well as access to undergraduate
and graduate students, many of whom are looking for research oppor-
tunities. They also have access to vast stores of knowledge through
their extensive hard copy and digital library systems. Zoo–university
collaborations have a long history, even going back to the time of
Charles Darwin, who questioned keepers and made observations of
animals at the London Zoo for his book The Expression of Emotions
in Man and Animals published in 1872 (Wemmer & Thompson,
1995).
A successful zoo–university partnership must be a two-way
street, with both partners deriving benefits. For example, zoos offer
incredible opportunities to train students in animal behavior research
methodology (Beck, 1974; Crockett, 1977; Rumbaugh, 1971). The
University of Washington in Seattle, for example, has taught a
“Behavioral Studies of Zoo Animals” course since the late 1970s in
cooperation with the Woodland Park Zoological Gardens (Crockett,
1977). This has not only resulted in numerous student publications,
but also helped to refine observational research methodology (Crock-
ett & Ha, 2010). Similarly, Disney’s Animal Kingdom theme park has
several scientists on staff and developed a cooperative relationship
with several local universities, including the University of Central
Florida. This includes adjunct academic appointments for Disney
scientists and the establishment of a Conservation Biology program
in which Disney scientists help to teach relevant courses (Lawson
et al., 2008).
The design of such cooperative programs, often formalized
through memoranda of understanding, is critical (Eisenberg, 1975;
Hutchins, 1988; Kleiman, 1996). For example, among other things,
injuries to students or faculty while on zoo grounds is a potential legal
and financial liability for zoos, which must be addressed in an agree-
ment. Conversely, it is also critical that the zoo is recognized for

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providing facilities, subjects, or personnel in any publications that

result and also has access to the results (Eisenberg, 1977; Hutchins,
1988; Kleiman, 1996). Unless properly managed, outside researchers
can also interfere with zoo operations, including animal management
and keeper routines. Involvement of zoo curatorial and keeper staff is
also important for promoting within-institution acceptance and
understanding of the relevance of research to the zoo’s core missions
of animal care, welfare, education, and conservation (Hutchins, 1988;
Lawson et al., 2008).

In-House Scientific Staff

Eisenberg (1975) suggested that an effective zoo research program is
best developed by having a permanently funded, separate department
within the zoo to manage it, and we agree. The presence of trained
scientists on staff indicates a strong institutional commitment and
works to fully integrate science into the day-to-day management of
the animal collection. Unfortunately, in a recent study of factors
facilitating zoo research, Anderson et al. (2010) found that a majority
of zoo-based research was being conducted by personnel without
advanced academic degrees. Furthermore, most research was con-
ducted and coordinated by curators and managers rather than by
dedicated scientific staff. The authors concluded that many zoos and
aquariums still did not prioritize research. The authors also found that
support of the zoo director was a critical factor influencing the success
of any zoo research program. That being said, many zoos, particularly
larger and well-funded organizations, have dedicated research facil-
ities and personnel on staff.
Benirshke (1996) and others have brought up the need for
multi-disciplinary research teams to tackle problems in zoo animal
management, husbandry, and care. There is no doubt that more
minds/perspectives are better than one, so cooperation between sci-
entists should be encouraged. When these teams involve scientists
with different areas of expertise, this is when true innovation is
possible.

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   ,  . ,   

Welfare and Other Ethical Considerations

The conduct of science on both nonhuman and human animals
has important ethical considerations that must be addressed. Any
research that has the potential to harm animals or humans, or, in
the latter case, to infringe on their privacy, is cause for concern and
must be carefully considered. This has been a hot topic for both
laboratory and field researchers in the past few decades (e.g., Farns-
worth & Rosovsky, 1993; Rudacille, 2000), and it has important
implications for zoo-based researchers as well (Hutchins et al., 1995;
Kirkpatrick, 1996; Norton, Hutchins, Stevens, & Maple, 1995).
So, how does an institution decide on whether or not a particu-
lar area of research or research project is acceptable or not? Essen-
tially, the knowledge gained from a study must outweigh any
potential cost in terms of diminished animal welfare. In the USA at
least, weighing these factors is the job of an Institutional Animal Care
and Use Committee (Bieletski, 1996; Kohn, 1996) or a Human Sub-
jects Review Board (Gillespie & Melbert, 2015). Such committees,
required under the US Animal Welfare Act, consist of institutional
scientists and curators, an attending veterinarian, and typically one
scientifically literate outside representative.
Such committees are responsible to the zoo’s director/CEO,
review research proposals, and have final say over whether a project
is allowed to go forward or not. In all cases, it must be assumed that
the public and media will have access to this information and that the
institution will be able to justify its decision based on not only the
scientific knowledge gained, but also that the subjects’ welfare was
taken into careful consideration.

Publication of Zoo Research

Zoo-based research has been published in a wide range of mainstream
and specialized peer-reviewed books and journals, such as Science,
Animal Behaviour, Applied Animal Behaviour Science, Journal
of Wildlife Management, Conservation Biology, and many others,

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depending on topic and content. Examples of peer-reviewed journals

and books specialized to the zoo and aquarium field are the open
access Journal of Zoo and Aquarium Research in Europe (www.jzar
.org/jzar) and Zoo Biology in North America, published by Wiley
(https://onlinelibrary.wiley.com/journal/10982361). Also available
are Der Zoologishe Garten, published by Elsevier (www.journals.else
vier.com/der-zoologische-garten), International Zoo Yearbook, pub-
lished by Wiley for the Zoological Society of London (https://zslpubli
cations.onlinelibrary.wiley.com/loi/17481090), and Aquarium Sci-
ences and Conservation, published by Springer (https://link.springer
.com/journal/10507). Zoo veterinary research and case studies can be
found in the Journal of Zoo and Wildlife Medicine (http://zoowildlife
journal.com/?code=zvet-site), the official publication of the American
Association of Zoo Veterinarians, the European Association of Zoo
and Wildlife Veterinarians, and the American College of Zoological
Medicine. Results of zoo and aquarium research can also be found in
non-peer-reviewed proceedings of regional zoo and aquarium associ-
ation conferences, such as those published (now online) by the Asso-
ciation of Zoos and Aquariums in North America (www.aza.org/
conference-proceedings).

Sustaining Zoo Research Programs

Sustaining zoo research programs takes sufficient funds to support
scientific personnel and a place to house them, computers and other
equipment, travel to scientific meetings, publication costs, and so
forth. Most zoos have limited funds to direct toward research. How-
ever, Anderson et al. (2010) found that a majority of existing zoo
research programs were funded out of the general operating budget.
They further suggested that “endowments and funding by granting
agencies, private foundations, and universities should be explored to
support science in zoos more broadly and effectively” (p. 573). That
being said, many scientists working at zoos do apply for and receive
grants on occasion from government wildlife and granting agencies
and from zoo-based regional (e.g., the Conservation Grants Fund of the

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   ,  . ,   

Association of Zoos and Aquariums in North America) and institu-

tional granting programs set up for that purpose (Hutchins & Ballen-
tine, 2001; Lawson et al., 2008).


Zoos have come a long way in the development of their considerable
research potential, but the task is far from complete. This chapter has
documented the tremendous scope of scientific research conducted by
contemporary zoos, both ex situ and in situ. This volume aims to
illustrate the scope of zoo research by providing specific examples
illustrating its diversity and impact. Zoo studies have improved
animal care, husbandry, and welfare, improved education, and con-
tributed to our knowledge of the natural world and to wildlife conser-
vation. We also documented the fact that many zoos have established
organized research programs to facilitate research and sustain them
through operating budgets and grants. Besides publishing the results
of their research in mainstream scientific journals and books, zoos
have also established and supported their own specialized book series
and journals. Despite this incredible progress, there is still much to
be done before zoos can reach their full potential. Among these, we
would include the need:

(1) To establish scientific programs at more institutions, particularly smaller

and medium-sized institutions, and hire dedicated and well-trained
scientists to run them;
(2) To establish more off-site facilities dedicated to animal research, which
can offer better opportunities for larger sample sizes and controlled
studies;
(3) To continue to pursue viable partnerships with universities, non-profit
conservation organizations, and government wildlife and research agencies
that have relevant scientific and conservation expertise;
(4) To expand research efforts to include a wider range of taxa, including
reptiles, amphibians, small mammals, insects, and other aquatic and
terrestrial invertebrates that are currently underrepresented in zoo-based
science;

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(5) To establish regional research priorities through gap analyses that assess
what is currently known and what needs to be known to solve particular
challenges in zoo and wild animal care, husbandry, welfare, health, and
conservation;
(6) To expand the role of zoos in field (in situ) research into animal behavior,
biology, and ecology and successful conservation methods;
(7) To continue educational research designed to improve the impacts of
zoos on visitor behavior – located in urban areas, zoos may be an ideal
place to promote behavior change, leading to wildlife-friendly practices
at home and in the global community;
(8) To promote multidisciplinary study teams either within or between
cooperating institutions or between regional zoo associations that can
both increase sample size and tackle “big picture” challenges that are
common to all;
(9) To diversify the ways in which zoo research is funded through the raising
of endowments and the pursuit of grants and donations from private
and corporate funders; and
(10) To better publicize the results of zoo-based research both to build public
support and to increase understanding of the role of zoos as research
facilities.

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