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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
Comprehensive reference detailing behavioral assessment techniques, behavioral approaches, and practical tools for animal welfare across different species
A Guide to Managing Zoo Animal Welfare delivers a step-by-step guide to behavioral assessment approaches, techniques, and tools for animal welfare with an emphasis on animals living in zoos and aquaria. The authors develop a unique “balance-based” approach that can be used to assess and enhance the welfare of a diverse range of species. Backed by extensive scientific literature, this book also provides foundational context to help readers to understand why the authors give these recommendations and guidelines.
This book is divided into three sections. Section I details background concepts and goals, discussing the animal mind through neuroscience, psychology, and behavior, even questioning wild animal behavior’s validity as a template for captive animal behavior. Section II details the core behavioral Needs of animals, investigating, acquiring rewards, and exerting control. Section III explains how to practically assess if animals’ Needs are met and address deficiencies, covering topics like food, space, and social rewards and methods to make environments dynamic.
A Guide to Managing Zoo Animal Welfare includes information on:
- Proximate and ultimate reasoning for the existence of each of the Needs
- Welfare benefits of meeting the Needs, including positive affect, maintaining homeostasis, passing on genes, and learning through reinforcement
- The highly variable personalities of individual animals and different animal species, as well as why personality is an important facet of animal welfare
- A guide for assessing animal welfare using the Three Needs model
Presenting highly unique insight and a novel approach, A Guide to Managing Zoo Animal Welfare is an essential learning resource for professional animal behaviorists in zoos and aquaria, researchers in animal behavior and animal welfare, and students in veterinary sciences and zoology programs of study.
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Seitenzahl: 413
Veröffentlichungsjahr: 2025
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Table of Contents
Cover
Table of Contents
Title Page
Copyright Page
Dedication Page
1 Background Concepts and Goals
Keeping Animals
Affective Neuroscience
Motivation Psychology
Animal Behavior
References
2 Is Wild Animal Behavior a Proper Template for Captive Animal Behavior?
References
3 Investigating
Proximate Versus Ultimate Drivers of Behavior – A Note on Terminology
Investigation
Why Investigate?
Proximate Drivers of Investigation
Animals Under Human Care Retain Their Drive for Understanding
References
4 Acquiring Reward
Why Acquire Rewards?
Proximate Drivers of Reward Acquisition
Acquiring Rewards Supports Welfare of Animals in Human Care
References
5 Exerting Control
Why Exert Control?
Proximate Drivers of Exerting Control
Opportunities to Exert Control for Supporting Well‐Being in Captive Animals
References
6 Balance: Good Welfare Through Psychological Resilience
References
7 Animal Personality: For Every Animal, All Needs Are Important, but They Are Not Equal
References
8 What It Looks Like When the Needs Are Met and Not Met
Meeting Needs
References
9 Assessment. Asking Animals, “Are Your Needs Met?”
The Balance Sheet Approach Using Direct Observation
Developing Observation Methods
Developing a List of Behaviors to Look for
Assessing the Data
The Balance Sheet Approach – Using Caretaker Interviews
Summarizing the Interview
Direct Tests and Probes for Mood‐Like State
Anticipatory Behavior
Cognitive Bias
Novel Objects
Assessing the Opportunities
References
10 A Practitioner’s Guide to Meeting the Needs
Supporting Investigation
Supporting the Acquisition of Rewards
Supporting Exerting Control
Availability of Opportunities
Relevance of Opportunities
Ability to Engage
Surrogate Behaviors, Surrogate Behavioral Sequences, and Cross‐Domain Carryover of Opportunity Effects
References
11 Exceptional Cases: Geriatric, Chronically and Acutely Ill, and Injured Animals
Does the Animal Experience Themself as Physically Healthy?
Does the Animal Experience Positive Psychological States?
Individual Variation and Ability to Experience Positive Welfare
Does the Animal Experience Independence?
Does the Animal Experience Appropriate Social Interactions?
Does the Animal Experience a Complex, Appropriate Environment?
Balance Among Domains
Applying the Balance Model to Assess Exceptional Cases
Change Over Time
References
12 Scenarios
Scenario 1: Rock Hyrax
Scenario 2: Parrot
Scenario 3: Rhino
Scenario 4: Guinea Baboon One
Scenario 5: Guinea Baboon Two
Scenario 6: African Painted Dog
Scenario 7: Reticulated Python
Data Collection – General
Data Collection
Data Interpretation and Suggested Management Changes
References
Index
End User License Agreement
List of Tables
Chapter 2
Table 2.1 The three most used definitions of animal welfare. Each definitio...
Chapter 8
Table 8.1 Example generic behaviors that reflect whether a Need is being me...
Chapter 11
Table 11.1 (A) Behavioral observations and their corresponding positive or ...
Chapter 12
Table 12.1 Rock Hyrax ethogram.
Table 12.2 Rock hyrax ethogram with the three Needs model, valence of the b...
Table 12.3 Parrot ethogram with definition of behaviors and types of observ...
Table 12.4 Parrot ethogram and three Needs related to each behavior.
Table 12.5 Modifier for parrot ethogram – to be used for perch behaviors an...
Table 12.6 Black rhino ethogram with behavior definitions and type of sugge...
Table 12.7 Black rhino ethogram with the three Needs related to the behavio...
Table 12.8 Guinea baboon ethogram – for scenarios 4 and 5.
Table 12.9 Guinea baboon ethogram and three Needs to which the behaviors re...
Table 12.10 Special all occurrence behavior with additional note for juveni...
Table 12.11 Ethogram for geriatric African painted dog.
Table 12.12 Ethogram for geriatric African painted dog and three Needs rela...
List of Illustrations
Chapter 2
Figure 2.1 Two views of welfare state. The natural state perspective assumes...
Figure 2.2 Historically, zoos have gone through many changes in focus and mi...
Chapter 3
Figure 3.1 An illustration of proximate and ultimate causes in animal behavi...
Figure 3.2 Animals exist within a dynamic world. To survive, animals must be...
Figure 3.3 (a) Example of a standard environment compared to a dynamic one. ...
Figure 3.4 Predicted net welfare impacts of animal responses to reliable and...
Figure 3.5 Traditional visitor effect (Hosey’s) describes zoogoers as having...
Chapter 4
Figure 4.1 Examples of behavioral chains. (a) Crane mating behaviors: identi...
Figure 4.2 The model of anticipatory behavior that Watters modified from van...
Chapter 5
Figure 5.1 Relationship between environmental predictability and animals’ se...
Chapter 6
Figure 6.1 A valence/arousal graph. These graphs often place emotional state...
Figure 6.2 A resilience model. Animals experiencing a similar frequency of p...
Chapter 7
Figure 7.1 Expectations: for the expression of anticipatory behavior depende...
Chapter 9
Figure 9.1 Accurate interpretation is important when using behavior as a dia...
Figure 9.2 (a) Traditional time budget presentation calculated from observat...
Figure 9.3 Examples of daily, weekly, and monthly balance sheets for two ani...
Figure 9.4 Decision tree to assist picking an assessment plan.
Figure 9.5 Steps that should be taken when using anticipatory behavior (AB) ...
Chapter 10
Figure 10.1 A belt feeder such as this one from Pentair, used by Watters et ...
Figure 10.2 Feedpods are automatic feeders that can be both scheduled and dr...
Figure 10.3 Primate Canopy Trails at St. Louis Zoo allows animals to choose ...
Chapter 11
Figure 11.1 Example plots of domain scores over time. In this example, a con...
Chapter 12
Figure 12.1 Three Needs graph for male rock hyrax.
Figure 12.2 Three Needs graph for blue and gold macaw.
Figure 12.3 Three Needs graph for female black rhino.
Figure 12.4 Three Needs graph for juvenile male guinea baboon.
Figure 12.5 Three Needs graph for adult male guinea baboon.
Figure 12.6 Three Needs graph for aged male African painted dog.
Figure 12.7 Three Needs graph for reticulated python.
Guide
Cover Page
Title Page
Copyright Page
Dedication Page
Table of Contents
Begin Reading
Index
WILEY END USER LICENSE AGREEMENT
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A Guide to Managing Zoo Animal Welfare
A Behavioral Approach
Dr. Jason V. Watters
Adjunct Associate Professor
Animal Science
University of California
Davis, California
USA
Dr. Bethany L. Krebs
Executive Editor of Zoo Biology
San Francisco, California
USA
This edition first published 2025© 2025 John Wiley & Sons Ltd
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For every beast whether they have four legs, two, none, or many:
fins, wings, arms, tentacles, or antennae.
1Background Concepts and Goals
Keeping Animals
For many reasons, people manage numerous species of animals in what we most frequently refer to as captive conditions. Now, more than ever before, there is a major emphasis on the psychological well‐being of the animals cared for in any industry of which they are a part. In this book, it will seem as if we make a distinction between wild or exotic captive animals and domestic animals and focus our examples on the former. This is not to say that the concepts developed here are not also applicable to domestic animals. Our focus on exotics is aimed at the development of a general approach. We develop this approach for two primary reasons. One reason is that the environments and behavioral outcomes associated with them that non‐domestic animals live in under human care are a traditional focus of inquiry when it comes to assessing the psychological state of these animals. Less frequently do we compare the domestic animal’s environment with their wild counterpart’s. Such comparisons are not realistic because, due to artificial selection for domestication, there is no naturally selected wild counterpart to a derived domestic animal. Who is to say that the behavior of the feral mustang is the standard to which the barnyard stallion should be held? A dog is not the same animal as a wolf. Instead, but for subtly different reasons, we compare domestic animals to what we know of their wild ancestors – who are most commonly extinct. The dog and the wolf may have shared an ancestor in the form of some ancient extinct wolf, but they have since gone their separate ways. Nevertheless, an understanding of the evolutionary history of the dog and the wolf is likely to support, not guarantee, caring for either animal in the modern world – regardless of the environment we find them in. Thus, by focusing on wild, exotic animals, we hope to emphasize the point that the psychological care of animals is not dependent on the exact replication of the environmental conditions under which the animals live in the wild. We also hope to emphasize that it is possible to develop a general approach to thinking about and supporting animals’ psychological well‐being that can be used to care for individuals of numerous species, wild and domestic.
As a result of evolution’s slow progression of tinkering on a core blueprint, there are fundamental similarities across species that support the application of a general approach to promoting animal welfare. Most of us are aware of some of these similarities, such as the generalized vertebrate body plan, but they also include subcortical emotional circuitry that is also widespread in the animal kingdom. The Cambridge Declaration on Consciousness (2012) resulted from a gathering of prominent neuroscientists with varied specializations in anatomy, physiology, pharmacology, and cognition. In developing the Declaration, these experts considered the convergence of evidence that indicates that nonhuman animals, both vertebrate and invertebrate, possess the neuroanatomy that generates and processes affective states. Experimental evidence shows that when these subcortical brain structures are stimulated, behaviors expressed are like those expressed when humans or other animals experience rewarding or punishing affective situations. The Declaration indicates that these emotional brain regions are present in all mammals and birds and that the neural circuits necessary to promote attentiveness, sleep, and decision‐making arose long enough ago that they are also present in insects and cephalopod mollusks.
The Declaration states that “emotional feelings” arise from a set of subcortical brain regions that are similar across species and that there is adequate evidence to suggest that humans and nonhuman animals possess “evolutionarily shared primal affective qualia.” This is to say that intrinsic self‐assessment of positive and negative feelings is evolutionarily old. All animals do this. While the Declaration makes no statement regarding reptiles, amphibians, or fish, the inclusion of insects, cephalopods, birds, and mammals suggests that these taxa would also be included in the thesis of the Declaration were there adequate data to provide examples. Today, evidence is starting to arise that suggests this is the case. This support comes from studies that demonstrate phenomena such as cognitive bias in bees (Bateson et al., 2011) and anticipatory behavior in fish (Reebs and Lague, 2000, Galhardo et al., 2011) and other species (Anderson et al., 2015, McGrath et al., 2016, Clegg et al., 2018, Podturkin et al., 2023). Animal welfare scientists consider these behavioral responses to reflect animals’ own perception of their affective state (Moe et al., 2006, Van der Harst and Spruijt, 2007).
A description of the core mind here may help to understand some of the justification and techniques described in the chapters that follow. A mind is the cumulative set of traits that enable awareness, decision‐making, memory, and affective response (i.e. internal reflection). Overall – the mind, like other traits, serves a developed function – and that is to assess and relate to the individual information regarding their own state as well as their potential state given their perceived circumstances. An individual’s mind determines the interpretation of circumstances, both cumulative and present, and generates from this determination the individual’s perception of balance. Balance in this way refers to the aggregate assessment of the internal outcomes associated with events the individual experiences. Other authors have referred to this state as mood‐like state (Yang et al., 2013, Mendl and Paul, 2020, Bliss‐Moreau and Rudebeck, 2021). The cumulative emotional experience of an animal builds their mood‐like state (Kremer et al., 2020, Mendl and Paul, 2020). As we hope to apply this thinking to support animal welfare, we use the term “balance” to describe our approach to measuring and managing this state. Experiences are either affectively positive, negative, or neutral (Gasper et al., 2019, Gasper, 2023). Positive and negatively valenced affect can be driven by either tangible outcomes – as in the cases of procuring food or being injured – or intangible ones – as in the cases of information acquisition or becoming fearful. In either case, tangible or intangible outcomes, there is an affective response – essentially, and very simply put, the individual either feels good about the outcome or does not feel good about it. Neutral affective outcomes result in what would appear to be behavioral indifference or a lack of preference (Gasper et al., 2019, Gasper, 2023). The mind thus serves to evaluate these affective outcomes and determine whether the animal should attempt to promote further similar events or avoid them. Thus, the mind serves the function of monitoring and reporting on affective outcomes. At the very least, a balance between positive and negative outcomes ensures that this core mind remains healthy. In this sense, the mind requires specific nurturing and can be unhealthy even when the body appears healthy. While we acknowledge that the brains of many species have evolved to facilitate advanced cognitive abilities, problem‐solving, and even self‐recognition (Roth, 2015, Mitchell, 2016, Gallup Jr and Anderson, 2020), we assert that the core mind is the seat of fundamental emotion, and that affect is a driving force in supporting evolutionary fitness.
Minds, like bodies, need to be nurtured. When an animal does not consistently live in a state where they achieve that balance between positive and negative outcomes, they can be in a state of negative welfare. This negative state is psychologically damaging (Désiré et al., 2002). The primary goal of this book is to provide a general technique to assess animals’ own perspectives of their welfare state and to suggest techniques to provide positive opportunities when an animal is negatively out of balance. Throughout the book, we will refer to animals’ core behavioral Needs, investigating, acquiring rewards, and managing one’s own processes. These core Needs are evolutionarily old and the logic that we will lay out in this book suggests that ensuring that the Needs are met provides for a balanced state of welfare. Our approach is thus a focus on the fifth domain – mental state – of the five domains model of animal welfare (Mellor and Beausoleil, 2015). Throughout, we will capitalize the word Need when we refer to these three Needs as a means to set these Needs apart from other needs that animals may have.
Our concept of animal welfare is a kingdom‐level individual property. It occurs across phyla and is not a species‐level property. There is a reasonable amount of evidence supporting the notion that the three motivational Needs described herein are shared across species and that they are thus of fundamental importance at the level of the kingdom. Phenomena such as cognitive bias in species ranging from bees to bears, anticipatory behavior in species as disparate as rhinoceroses and fish, contra‐freeloading of numerous species of birds and mammals, preferences for enriched spaces from turtles to rats, and widespread reward seeking behaviors support the notion that individuals across the animal kingdom share the motivational Needs we describe. This understanding is core to the cross‐species applicability of our approach (Pacheco et al in prep). Our approach centers on assessing the three Needs in each animal who draws our focus. We specifically attempt to determine if the animal is showing us, with their behavior, if they are able to meet the three Needs. However, because the mechanism by which each animal interfaces with the environment differs, we seek to understand species typical behaviors that reflect engaging the motivations that underly the three Needs. We often ask, “What does investigatory behavior look like in the species we are currently observing?”
Applied animal welfare workers often approach adjusting animals’ mental state through the application of environmental or behavioral enrichment (Brydges et al., 2011, Douglas et al., 2012). However, this book is not aimed at being a clearinghouse for animal enrichment ideas. Many of those are already available. In its simplest sense, an animal’s mind is nurtured when the animal acquires meaningful and sometimes novel information, occasional and predictable success and has behavioral options that permit them to express agency. This novel information may be related to understanding one’s environment or understanding the consequences of one’s actions. The following chapters will elaborate on topics related to understanding what minds need and how to give it to them. Part of the book is written as a handbook for those caring for numerous species of animals in zoos, aquariums, and sanctuaries. Thus, while the primary concepts are general, there are specific sections pointed directly at this intended audience. These sections aim to further convince this audience that the psychological well‐being of the individual animal is of utmost importance to meeting their institutional goals. Throughout the book, we will attempt to provide necessary background but not excessive details in the hope that this will facilitate quick understanding of the key points while not dwelling on minutia and specific details of the supporting research. The goal, after all, is to support the practice of caring for animals’ minds, not to justify it.
The argument for the core behavioral Needs is developed here with consideration of the findings of three primary research disciplines. The disciplines use differing and sometimes overlapping techniques to evaluate behavior, its drivers, and its outcomes. Where techniques overlap, they help to demonstrate generality and where techniques are novel to a discipline, they help to validate findings. Some techniques are not practiced across fields because of their invasiveness or varied feasibility with certain subjects. For example, human psychology researchers can hold conversations with their subjects while animal behavior researchers cannot. Similarly, affective neuroscientists can stimulate specific brain regions and observe responses.
Several disciplines separately indicate similar factors that lead to positive psychological and behavioral outcomes for both animals and people. Taken together, these disciplines suggest a simple set of core behavioral Needs. Specifically, these disciplines are Affective Neuroscience, Motivation Psychology, and Animal Behavior. In essence, we use these disciplines’ emphasis on similar goals to build support for our approach through a triangulation wherein we look for the overlap of findings across fields or the application of those from one field to another. Jaak Panksepp is our inspiration for applying this approach.
In the end, we hope to provide a way to observe and measure animal behavior that is informed by these disciplines and that helps the observer to understand if animals’ core Needs are met. Another goal of the book is to suggest a general technique that can be useful for monitoring and improving animal welfare and is not reliant on expensive technology or testing. Its efficacy is observable through animals’ behavior. In the sections that describe animals’ Needs, we will present evidence from each of these areas in support of that Need’s importance.
Affective Neuroscience
Affective neuroscientists focus their efforts on understanding the neurological mechanisms of emotion. They often use techniques that directly measure the neural responses to specific stimuli. For example, Jaak Panksepp, whose work demonstrated seven basic mammalian emotions used a combination of neuro‐stimulation and recording techniques coupled with various stimuli to track the seat of several emotions in the brain as well as the situations that stimulate them and the behavioral responses associated with them (Panksepp, 2005a, 2011). We follow Panksepp in using a triangulation approach. He included evidence from behavioral, psychological, and neuroscientific studies to elucidate what he called core affective states (Panksepp, 2005b). Specifically, Panksepp was a pioneer in the field of affective neuroscience, demonstrating that raw affects arise from the sub‐neocortical neurophysiologies of emotional action systems in the mammalian brain. These findings were in stark contrast to the assumptions of behavioral neuroscience and cognitive neuroscience that placed the development of affect as a higher order brain process (Panksepp, 2005b). As a result of these disparities across neuroscience fields, Panksepp believed that affect was overlooked in studies of basic animal biology. Though affect is becoming a central paradigm of animal welfare research.
Panksepp and colleagues’ work lays a foundation for understanding that there are evolutionarily old behavioral Needs. His findings suggest that the core emotions are subcortical and not the result of higher order cognitive function. It is this work that demonstrates the locations in the brain where stimuli generate affective responses and associated behaviors that suggest that mind and emotion do not require higher order cognitive capability. In addition, work in this field suggests that when certain basic emotions are not experienced, psychological welfare is compromised (Alcaro and Panksepp, 2011).
Motivation Psychology
Human psychologists are interested in how the state of a person’s mind drives their behavior. Scientists studying humans can interview their subjects and ask informed questions. This process, where the subject of study describes to the researcher their feelings and actions, is referred to as self‐reporting (Paulhus and Vazire, 2007). Those who work on understanding the connections between emotional states and behavior advance these understandings with the self‐report process. While the credibility of self‐reported data is sometimes criticized (Paulhus and Vazire, 2007), the self‐report procedure is perhaps the sole technique that allows the direct coupling of affective state with behavior. Given the possibility of widespread affect‐driven behavior as proposed in the Cambridge Declaration on Consciousness, it seems possible that educated observers could use animals’ behavior as a sort of a self‐report and a means to assess animals’ emotional state.
E. Tory Higgins and colleagues have developed (Higgins, 1997) the conceptual framework of “effectiveness” that describes motivations aimed at generating positive emotional outcomes. The theory of effectiveness describes three primary motivational goals that support positive well‐being. Higgins and colleagues refer to these motivational goals as truth, value, and control. Their research demonstrates that supporting these motivations leads to positive well‐being. Becca Franks has extended Higgins’ approach to the field of animal welfare (Franks and Higgins, 2012). She has demonstrated that the concept of effectiveness carries over to animals.
Animal Behavior
Animal behavior research has expanded humans’ understanding of numerous factors including animal movement, communication, mate choice and sexual selection, foraging decisions, predator–prey interactions, and many more areas. Much animal welfare research is housed within this discipline. Here, specific questions regarding what animals want, one of Marian Stamp Dawkins (Dawkins, 2021) two components of animal welfare (the other being health), are often addressed. Additionally, animal behavior researchers are now frequently asking questions about how animals feel – in part through approaches that draw on other fields but rely on behavioral observation. These two lines of inquiry, what animals want and how they feel, are tools for assessing animals’ mental health.
Studies that investigate animals’ preferences, anticipatory responses, decision‐making, willingness to work, and response to novelty are among those that point to animals’ core behavioral Needs. They reinforce our understanding of animals’ basic motivations that we believe must be supported to promote animal welfare.
We acknowledge that there are many interpretations of the term animal welfare (Fisher, 2009, Watters et al., 2021). Its definitions vary and, as a result, we feel it is important for those writing on the subject to define it so that their meaning is not misinterpreted by others who are interested in the subject but perhaps ascribe a different meaning to it. Indeed, as animal welfare research expands and takes on more nuanced forms of investigation, this simple description upon planning research or describing it will help workers to justify their choice of variables to measure and the techniques they use to assess them.
In previous work, we have described animal welfare as being a state of balance between animals’ positive and negative affective experiences (Watters et al., 2019). We have suggested that animal welfare is the animal’s own perception of their psychological state (Watters and Krebs, 2019). We have assessed this reflection with various behaviors and have justified doing so with chains of logic. We consider an animal’s welfare to be their cumulative emotional state. Often referred to as an animal’s mood‐like state (Mendl and Paul, 2020), this cumulative state reflects the overall balance between positive and negative emotional outcomes. Some circumstances lead to positive affective responses while others result in negative ones. It is the state of balance between these positive and negative emotional outcomes that we measure when we ascribe an animal’s welfare state. We admit that we do not know exactly the duration that we should assess to determine a cumulative welfare state and suggest that this duration may be dependent upon the presence of very strongly valenced, highly aroused affective outcomes. For example, experiences that produce very strong fear responses are likely to cause shifts to a generally negative affective state that is relatively long lasting compared to episodes that result in brief moments of fear followed by recovery (Jones and Boissy, 2011). We also suggest that because they are built on cumulative outcomes, mood‐like states should be observable repeatedly over relatively short time frames and can persist for very long ones, though our key premise is that they can change.
We will provide suggestions for developing protocols aimed at observing animals’ cumulative affective state. These protocols will help observers determine the duration of observation necessary to draw conclusions about animals’ own perceptions of their well‐being. The approach we will describe combines observation across time with specific situational observation as there are situationally expressed behaviors that reflect some degree of this cumulative state.
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2Is Wild Animal Behavior a Proper Template for Captive Animal Behavior?
Most animal welfare workers acknowledge three types of definitions of animal welfare: natural state, biological function, and affective state. The three definitions differ in philosophical leanings and appear to be applied by workers in different fields. Traditionally, natural state definitions have been valued in zoos and by the public, biological function by veterinarians, and affective state by psychology‐leaning academics.
Natural state definitions of animal welfare compare animals under human care to wild animals. They assume that in the wild, animals have all the factors that support positive welfare. Those who ascribe to natural state definitions of animal welfare place high value on the behaviors that are observed in wild animals as being indicators of positive welfare. Indeed, the founder of the field of zoo biology, Heini Hediger, once wrote, “the standard by which a zoo animal is judged should be the life that it lives in the wild.” Though Hediger himself did not interpret this statement literally when considering zoo animals, there can be a tendency for both zoo biologists and laypersons to do so. Indeed, there is a prominent emphasis on building natural looking enclosures for the display of zoo animals that is accompanied by an emphasis on attempting to ensure that the animals can “act” in a fashion like their counterparts in wild settings. While primarily an education tool, aimed at providing deeper information about animals’ natural history, their native environment and conservation status this approach to exhibitry supports a natural state philosophy of animal welfare.
Biological function definitions of welfare assert that animals’ well‐being is supported when they can maintain health and physiological function – including reproduction if possible. These are relatively early definitions of welfare and tend to focus on the minimization of suffering. Distressed animals can suffer physiological and health concerns. For example, Wielebnowski (2003) points out that chronically distressed animals may experience “immune deficiency, reproductive suppression, growth reduction, muscle wasting, gastrointestinal dysfunction and impaired brain function.” Many of these outcomes are easily measured by those trained in animal health, veterinarians being among the earliest animal welfare practitioners. An additional aspect of these types of definitions of welfare is that they provide guidance for the development of resource‐based assessments of welfare. For example, animals with specific physiological temperature requirements clearly must be kept within a certain range of temperatures to achieve proper biological function.
The third primary line to describe animal welfare refers to the animal’s affective state. Those assessing welfare with this approach are concerned with the animal’s emotional state. Their focus is to understand how to determine whether animals have psychologically positive or negative experiences and how they express their perception of them. In practice, emphasis is placed on ensuring that animals experience pleasurable moments and avoid negative experiences such as fear or anxiety. This approach thus is most concerned with animals’ own perceptions of their state, though it can be confounded by humans’ expectations of what animals should consider positive as well as well‐meaning (Watters et al., 2021) but misguided attempts to avoid any negative experiences at all. Temporary moments of discomfort are not typically reflective of overall poor welfare. Those applying an affective state view of welfare, however, may be freed from comparing an animal’s living environment to that in which their ancestors or relatives evolved. Instead, they can focus on reading individual animals’ own indications of their perceived state. Underlying this philosophy is the possibility that there are surrogate behaviors that, when expressed, are positive feeling for animals – even if they are not a regular component of a wild behavioral repertoire. For example, captive animals are often provided with puzzle feeders that evoke problem‐solving and often elicit behaviors that would not be a primary component of a wild behavioral repertoire. Although the exact behaviors required to retrieve food from a feeder may not correspond to wild behaviors, the goal‐motivated problem‐solving behaviors exhibited do and animals’ regular use of these opportunities suggest they experience positive moments in doing so (Table 2.1).
The three philosophies of animal welfare are both synergistic and antagonistic to each other. Presumably, a happy, healthy animal that can behave in a natural fashion is one in the most positive state of welfare. However, as we know from human psychology, there is a potential for physiologically unhealthy individuals to perceive that they are happy or vice versa (Carel, 2007). Indeed, it is possible that positive affective experiences support physiological health (Cohen and Pressman, 2006). With further study of affective experience, it is now clear that emotional responses are indeed biological functions in and of themselves. Thus, the continued development of welfare science is seeing a unification of the biological function and affective state frameworks (Hemsworth et al., 2015). It is also possible for natural occurrences to include threatening or uncomfortable ones and that these can persist for long durations – leading to apparent tension between natural state and alternative frameworks of animal welfare. For example, it would seem inappropriate to assign good welfare to a wild prey animal who has been separated from their social group and is wandering across a landscape wrought with predators in search of haven, food, and social partners although this is a common circumstance for many wild animals.
Table 2.1 The three most used definitions of animal welfare. Each definition has a different standard for determining when interventions are needed to improve animal welfare.
Common Definitions of Animal Welfare
Physical Health and Function
Natural State
Affective State
How welfare is defined
Animals whose bodies are healthy and functioning normally are experiencing good welfare.
Animals who can express the same behaviors as their wild counterparts are experiencing good welfare.
Animals who experience positive psychological feelings are experiencing good welfare.
When interventions are needed
Interventions applied when signs of disease or changes to physical function appear.
Interventions applied when deviations from behavioral patterns of wild counterparts appear.
Interventions applied when animals are experiencing negative psychological feelings.
We suggest that this tension is also unnecessary and that further unification of welfare frameworks is possible. First, though, it is somewhat useful to tear down the philosophy that captive animals that do not express behaviors as their wild counterparts do experience negative welfare. Wild animal behavior is often used as a reference for the behavior of captive animals – even though their environments are almost always fundamentally different. By comparison, numerous studies that investigate the behavior of wild animals of the same species in different populations find considerable differences in numerous behaviors across populations. These differences are noted in behaviors from foraging to group structure to reproductive tactics employed and many more. Indeed, the experimental currency of the animal behavior scientist is exactly the ability to vary behavior with changes in animals’ external or internal states. Behavior varies widely with ecology, group structure, population density, resource availability, and a very long list of other factors. Thus, the zoo biologist striving to emulate natural wild animal behaviors in the captive setting must first ask, “Which wild?” Perhaps instead, they should ask, “Why bother?”
Do zoo cheetahs dream about chasing gazelles at 70 miles an hour? Do polar bears crave long walks on endless sea ice? Ultimately, the natural state philosophy is an evolutionary one. It is hard to argue against the logic of supporting animals based on their evolutionary needs. Indeed, there is something that feels naturally “right” about supporting animals’ evolved needs. It is perhaps because of this feeling that people have that the natural state philosophy of animal welfare seems to maintain prominence even when those who care for animals recognize the fundamental differences between wild and captive states. Captive environments differ from wild ones with respect to many components. They also have similarities. Nevertheless, the likelihood that captive animals should act in the same fashion as wild ones is perhaps even smaller than the expectation that two wild populations in markedly different habitats should maintain the same behavior, social structure, foraging techniques, etc. Yet, it is not uncommon to find recent publications – particularly in the area of animal welfare – with statements that indicate natural behaviors as indicators of positive animal welfare. Suggesting that a polar bear who lives in a zoo cannot have positive welfare because she cannot walk across hundreds of square miles of sea ice does not provide a solution for giving that bear positive welfare. There are no zoos that have animal facilities as large as the natural environment of even a single polar bear. So, are we to euthanize captive polar bears for their lack of space? No. It is not the case that every polar bear living in a zoo suffers poor welfare. Thus, there is some fallacy in the assumption that the observed natural state is the default best state. Instead, we suggest that the observed state is the expression of the interaction between underlying evolved motivations and current and past circumstances experienced by the individual.
Wild polar bears don’t crave long walks on endless sea ice. They need them. Captive bears do not. Wild bears need these walks not to satisfy their emotional needs, but to satisfy their physical ones. The resources that wild polar bears need to survive are widespread and not predictable (Ferguson et al., 1999). Indeed, it is likely the case for nearly any animal with a large home range that their resources require them to search large areas (Gittleman and Harvey, 1982, Ross et al., 2012). Yet, it seems easy to commit the logical fallacy known as “appeal to nature” when referring to the perceived welfare of animals in captive situations. This fallacy extends from the enclosures in which animals live to the behavior of the animals themselves. Indeed, extensive programs exist to support naturalistic behavior. As mentioned earlier, these programs developed from an education emphasis, and they include placing significant focus on animal exhibitry that looks like natural settings. They are often messaged as supporting “natural behavior.” It is here, in the architecture of zoo settings and the quest for natural behavior that the appeal to nature fallacy is most dangerously employed. This danger looms for animals housed by people who value the esthetics of place over animals’ emotional needs. This misappropriation of value likely happens unwittingly and because of zealous interpretation of the natural state definition of welfare. Instead, perhaps rather than looking for behaviors that appear as they do in the wild, the welfare‐minded animal care professional should look for behaviors that indicate positive welfare – be they natural in frequency and appearance or not (Figure 2.1).
Maple and colleagues drew on the work of environmental psychologist Robert Sommer who had studied prisons and referred to their design as hard architecture (Sommer, 1974) to point out that zoos frequently employed similar design and that “softening” exhibit spaces supported positive welfare (Maple and Finlay, 1987). Sommer suggested that hard architecture resulted in negative psychological outcomes for those who inhabited it, and Maple and colleagues carried this line of thinking over to zoos. The result of this and other efforts was a noble push toward larger, lusher enclosures for zoo animals. Larger spaces alone will not ensure positive welfare for zoo animals. Neither will soft substrates, real vegetation, nor physiologically neutral temperatures. Of course, it would be ludicrous to suggest that modern animal enclosures should not include these items but the notion that they and other static aspects of the environment are, in and of themselves, a substrate for positive welfare is also ludicrous. If the fundamental problem of zoo animal behavior is not addressed with exhibit design or animal management, then the problem will persist. That problem is that captive animal environments are profoundly not dynamic. Animals who live in these environments quickly learn that the resources they desire most are associated with cues that come from outside of these environments and often at predictable times. Thus, the exhibit becomes little more than a waiting room. Of course, this problem can be overcome with creative application of technology (though it rarely is).
