Dynamics and energetics of bottlenose dolphin (Tursiops truncatus) fluke-and-glide gait.

Intermittent locomotion composed of periods of active flapping/stroking followed by inactive gliding has been observed with species that inhabit both aerial and marine environments. However, studies on the energetic benefits of a fluke-and-glide (FG) gait during horizontal locomotion are limited for dolphins. This work presents a physics-based model of FG gait and an analysis of the associated cost of transport for bottlenose dolphins (Tursiops truncatus). New gliding drag coefficients for the model were estimated using measured data from free-swimming bottlenose dolphins. The data-driven approach used kinematic measurement from 84 h of biologging tag data collected from three animals to estimate the coefficients. A set of 532 qualifying gliding events were automatically extracted for estimation of the gliding drag coefficient. Next, data from 783 FG bouts were parameterized and used with the model-based dynamic analysis to investigate the cost benefits of FG gait. Experimental results indicate that FG gait was preferred at speeds of ∼2.2-2.7 m s-1. Observed FG bouts had an average duty factor of 0.45 and a gliding duration of 5 s. The average associated metabolic cost of transport (COT) and mechanical cost of transport (MECOT) of FG gait are 2.53 and 0.35 J m-1 kg-1, respectively, at the preferred speeds. This corresponded to a respective 18.9% and 27.1% reduction in cost when compared with model predictions of continuous fluking gait at the same average bout speed. Average thrust was positively correlated with fluking frequency and amplitude as animals accelerated during the FG bouts, whereas fluking frequency and amplitude were negatively correlated for a given thrust range. These results suggest that FG gait enhances the horizontal swimming efficiency of bottlenose dolphins and provides new insights into the gait dynamics of these animals.

Preference assessments as a tool to evaluate environmental enrichment.

Environmental enrichment is an important tool utilized to improve animal welfare in zoological institutions through opportunity for mental and physical stimulation. Many past studies have focused on the impact enrichment has on animal behavior; however, none have conducted preference assessments on enrichment items to examine the relationship between animals' preferences and interaction with enrichment over a 24-hr period. Ten-minute free operant, paired-choice preference assessments were implemented in Study 1 to determine the enrichment preferences of African lions (N = 3). Following Study 1, Study 2 was conducted, which examined the behavior of African lions with enrichment items over the course of 30, 24-hr trials to evaluate the relationship between preferences established in Study 1 and long-term interaction with the enrichment. Generalized estimating equations revealed a statistically significant relationship between the percentage of time the enrichment was approached first and the average duration of interaction in Study 1 to the total duration of interaction and the percent of hours interacted with the item in Study 2. Additionally, the first 2 min of the preference data resulted in comparable statistically significant findings, demonstrating shorter preference assessments can produce similar results. The results support our prediction that preference assessments can be used to estimate the amount of interaction with enrichment over the course of time, with preferred items being interacted with more frequently and in longer duration. Information gained from this study suggests preference assessments can be a time and cost-effective tool to evaluate enrichment preference and predicted efficacy.

The impact of in-person and video-recorded animal experiences on zoo visitors' cognition, affect, empathic concern, and conservation intent.

Currently there are mixed results regarding the ability for media or more specifically video to increase a person's interest in conservation. However, there is a growing amount of evidence that in-person experiences at a zoo or aquarium can increase a person's interest in conservation. The goal of the current study was to examine the difference between an in-person experience viewing a polar bear training session and watching a video of the same experience on cognition, emotion, empathic concern, and conservation intent. A total of 124 Brookfield Zoo members were randomly assigned to one of three conditions. Condition 1 was an in-person 10 min (Live Animal) experience viewing a training session with a polar bear. Condition 2 participants (Video Animal) watched a video of the same experience from Condition 1 and Condition 3 (Control) listened to the audio from Condition 1 but only viewed an image of one of our animal care specialists. Results suggest that the live condition is associated with higher probability of answering questions correctly, having a positive emotional experience, having greater empathic concern for wild bears, and wanting to get involved in conservation when compared to the control. These impacts were not observed for the video condition suggesting that for this study, watching a video of a training session was not an effective tool for getting people involved in conservation. Future research is needed to better understand this important topic, but we now have further evidence of the importance of in-person zoo experiences.

Linking stress and immunity: Immunoglobulin A as a non-invasive physiological biomarker in animal welfare studies.

As the animal welfare community strives to empirically assess how care and management practices can help maintain or even enhance welfare, the development of tools for non-invasively measuring physiological biomarkers is essential. Of the suite of physiological biomarkers, Immunoglobulin A (IgA), particularly the secretory form (Secretory IgA or SIgA), is at the forefront because of its crucial role in mucosal immunity and links to physical health, stress, and overall psychological well-being. While interpretation of changes in SIgA concentrations on short time scales is complex, long-term SIgA patterns are consistent: conditions that create chronic stress lead to suppression of SIgA. In contrast, when welfare is enhanced, SIgA is predicted to stabilize at higher concentrations. In this review, we examine how SIgA concentrations are reflective of both physiological stress and immune function. We then review the literature associating SIgA concentrations with various metrics of animal welfare and provide detailed methodological considerations for SIgA monitoring. Overall, our aim is to provide an in-depth discussion regarding the value of SIgA as physiological biomarker to studies aiming to understand the links between stress and immunity.

Viewing African and Asian elephants at accredited zoological institutions: Conservation intent and perceptions of animal welfare.

African and Asian elephants are popular within zoos, however there is currently limited information on how viewing them impacts zoo visitors. The goal of the current study was to examine the relationship between viewing elephants in zoos accredited by the Association of Zoos and Aquariums and zoo visitors' reported conservation intent and perceptions of animal welfare. Visitors were systematically selected to fill out questionnaires following elephant observation at nine facilities throughout North America. Questions included information on conservation predispositions, exhibit experience, exhibit perceptions, animal welfare perceptions, emotional experience, learning outcomes, conservation intent, and demographics. Results suggest that observing elephants engaged in a variety of species-typical behaviors and having an up-close experience was significantly correlated to visitors having a positive emotional response. The positive emotional response, combined with visitor conservation predisposition had a significant positive relationship with reported interest in getting involved in conservation. Perceptions of animal welfare were significantly related to a positive emotional experience driven by seeing animals engaged in a variety of active species-typical behaviors as well as exhibit perceptions and whether or not visitors thought it was important to have elephants in zoos. Exhibit perception was primarily correlated with exhibit size. The results provide factors that could help to increase visitor interest in conservation as well as the potential impact of viewing elephants in an accredited zoo. Facilities can use this information to help ensure their visitors have similar type experiences in order to inspire visitors' interest in conservation as well as positive perceptions of animal welfare.

Creating a common terminology for play behavior to increase cross-disciplinary research.

Historically, play behavior has been difficult to define. This likely stems from the number of different species, types of play, and context under which it occurs. In 2016, the Chicago Zoological Society - Brookfield Zoo hosted the Psychonomic Society leading edge workshop on the evolutionary and psychological significance of play. Sixteen experts attended from the diverse fields of African ethnology, animal behavior, animal science, animal welfare, cognitive psychology, cognitive zoology, comparative psychology, cultural anthropology, developmental psychology, educational psychology, ethology, neuroscience, primatology, and zoology. Approximately half of the participants studied human play and the other half studied non-human play. Before the workshop, participants were asked to send in either their personal definition of play or the one that they cite in peer-reviewed literature. Definitions were then reviewed to determine characteristics of play inclusive of all disciplines. The goal of the current study was not to do a literature review on play behavior, but to come up with a list of characteristics across all forms of play that could be used as a common terminology moving forward. Hopefully the results of this workshop and the current article will help to increase cross-disciplinary research in the field of play.

Zoo visitor perceptions, attitudes, and conservation intent after viewing African elephants at the San Diego Zoo Safari Park.

Elephants in the wild face several conservation issues. With the rebranding of zoos as conservation and education pioneers, they have the ability to both educate and inspire guests to action. The purpose of this research was to analyze visitor perceptions and attitudes toward elephant conservation and outcomes post-exhibit visit. A one-page survey was randomly administered to assess perceptions of elephant behavior, attitudes about elephant conservation, and intended conservation-related outcomes from September 2013 to January 2014. Principle component analysis identified three major components: concern for elephants in zoos, importance of elephants in the wild, and modification of nature. Visitors who scored highly on conservation intent were those with positive attitudes towards elephants in the wild and negative attitudes regarding the modification of nature. The greatest changes in conservation intent were a result of a self-reported up-close encounter and the ability to witness active behaviors. Providing guests with the opportunity to witness or experience such occurrences may aid in a more successful delivery of the zoo's conservation message. Further research into guest emotions and affective states in relation to viewing elephants in a zoological institution would provide greater insight into improving the guest experience and helping zoos meet their conservation mission. Zoo Biol. 35:355-361, 2016. © 2016 Wiley Periodicals, Inc.

Diurnal and nocturnal activity budgets of zoo elephants in an outdoor facility.

The present study examined the activity budgets of 15 African elephants (1 bull, 6 cows, 2 male juveniles, 2 female juveniles, and 4 male calves) living at the San Diego Zoo Safari Park during the summers of 2010 and 2011. Onsite behavioral data (n = 600 hr) were collected for approximately 12 weeks from 0400 to 0830 and 1100 to 2400 during the 2010 and 2011 summer season. Foraging was the most common behavior state during the day followed by resting, and walking. During the evening hours, the elephants spent majority of their time foraging, resting, and sleeping. The average rate of self-maintenance behavior events (dust, wallow, etc.) increased from 0600 to 0700, 1100 to 1500, and from 1700 to 1900. Positive social behavior events (touch other, play, etc.) remained high from 0500 to 2300, with peaks at 0600, 1300, 1500, and 1900. Negative social events occurred at low rates throughout the day and night, with peaks at 0600, 1900, and 2200. The majority of positive behavior events during the daylight and nighttime hours involved the mother-calf pairs. Furthermore, the calves and juveniles initiated approximately 60% of all social events during the daytime and 57% of all social interactions at night. The results of this study demonstrate the differences between diurnal and nocturnal activity budgets of a multi-age and sex elephant herd in a zoological facility, which highlights the importance of managing elephants to meet their 24 hr behavioral needs.

Utilizing vocalizations to gain insight into the affective states of non-human mammals.

This review discusses how welfare scientists can examine vocalizations to gain insight into the affective states of individual animals. In recent years, researchers working in professionally managed settings have recognized the value of monitoring the types, rates, and acoustic structures of calls, which may reflect various aspects of welfare. Fortunately, recent technological advances in the field of bioacoustics allow for vocal activity to be recorded with microphones, hydrophones, and animal-attached devices (e.g., collars), as well as automated call recognition. We consider how vocal behavior can be used as an indicator of affective state, with particular interest in the valence of emotions. While most studies have investigated vocal activity produced in negative contexts (e.g., experiencing pain, social isolation, environmental disturbances), we highlight vocalizations that express positive affective states. For instance, some species produce vocalizations while foraging, playing, engaging in grooming, or interacting affiliatively with conspecifics. This review provides an overview of the evidence that exists for the construct validity of vocal indicators of affective state in non-human mammals. Furthermore, we discuss non-invasive methods that can be utilized to investigate vocal behavior, as well as potential limitations to this line of research. In the future, welfare scientists should attempt to identify reliable, valid species-specific calls that reflect emotional valence, which may be possible by adopting a dimensional approach. The dimensional approach considers both arousal and valence by comparing vocalizations emitted in negative and positive contexts. Ultimately, acoustic activity can be tracked continuously to detect shifts in welfare status or to evaluate the impact of animal transfers, introductions, and changes to the husbandry routine or environment. We encourage welfare scientists to expand their welfare monitoring toolkits by combining vocal activity with other behavioral measures and physiological biomarkers.

Corrigendum: Utilizing vocalizations to gain insight into the affective states of non-human mammals.

[This corrects the article DOI: 10.3389/fvets.2024.1366933.].

Beyond the Five Freedoms: Animal Welfare at Modern Zoological Facilities.

The current manuscript highlights the aspects of an animal welfare program for a modern zoological facility. The program should be proactive to identify areas for continuous improvement as well as reactive to address any identified animal welfare concerns. The program should go beyond the five freedoms and utilize one of the more modern frameworks as a foundation for the program. The program should have an animal welfare committee where staff can submit animal welfare concerns without fear of retaliation. Ongoing monitoring of all individual animals should utilize both positive and negative indicators of welfare. Staff should be trained on the most current science and be able to understand key concepts about animal welfare. Facilities should also utilize new scientific findings to continuously improve animal care practices. Modern zoological institutions, including both zoos and aquariums, have an ethical responsibility to provide high levels of animal welfare for the animals under their professional care. Simply meeting minimum standards developed decades ago is not adequate, as animals should have the opportunity to thrive.

Correction: Miller et al. Behavioral Diversity as a Potential Indicator of Positive Animal Welfare. Animals 2020, 10, 1211.

In the original publication [...].

Integrating Reference Intervals into Chimpanzee Welfare Research.

Animal welfare researchers are committed to developing novel approaches to enhance the quality of life of chimpanzees living in professional care. To systematically monitor physical, mental, and emotional states, welfare scientists highlight the importance of integrating non-invasive, animal-based welfare indicators. This study aimed to create species-specific reference intervals for behavioral measures and physiological biomarkers. Specifically, we analyzed data from 40 adult chimpanzees (22 females, 18 males) residing at 16 zoological facilities to generate reference intervals for behavioral states and events, behavioral diversity, fecal glucocorticoid metabolites (GCMs), and fecal immunoglobulin-A (IgA). Comparisons of sex and age using linear regression models revealed significant differences for several behaviors. The proportion of time spent engaged in mutual/multiple social grooming significantly decreased as individuals aged. Furthermore, males spent a higher proportion of time performing aggressive contact behaviors and displaying to other chimpanzees when compared to females. Males also performed sexual examination behaviors at a higher rate than females. Behavioral diversity, fecal GCM, and fecal IgA did not vary by sex or age. In the future, values for individual chimpanzees can be compared to the ranges reported here for particular age/sex classes. Ultimately, animal care professionals can utilize reference intervals to make evidence-based decisions regarding management practices and environmental conditions.

Visitor reaction to pacing behavior: influence on the perception of animal care and interest in supporting zoological institutions.

Many publications within the field of zoo animal welfare have stated the importance of decreasing stereotypic behavior (e.g., pacing) to help ensure a positive visitor experience. The idea behind these statements is that visitors want to see animals engaged in natural behavior. Additionally, it is thought that watching an animal exhibit species-appropriate behavior could help increase a visitor's connection to wildlife and ultimately their interest in conservation. However, until recently, no information was available to validate such statements. The purpose of this research was to examine people's reaction to viewing an animal engaged in pacing behavior. Participants were randomly selected to fill out a survey after watching a short video of either a tiger pacing or resting (control). Results indicate that having viewed a tiger pacing significantly decreases people's perception of the level of care animals receive at that facility. In addition, people's interest in supporting zoos decreased as a result of viewing this behavior. Results are discussed from an animal welfare, business, and conservation perspective.

Environmental enrichment, training, and habitat characteristics of common bottlenose dolphins (Tursiops truncatus) and Indo-Pacific bottlenose dolphins (Tursiops aduncus).

In recent decades, animal welfare science has evolved to utilize a multidisciplinary approach to assess the welfare of animals in accredited zoos and aquariums. Science-based animal welfare assessments have become an essential component of management programs and widespread application is expected by animal care professionals. Management practices for bottlenose dolphins in accredited facilities incorporate several programs that potentially impact animal welfare including environmental enrichment and animal training. Additionally, habitat characteristics, such as the dimensions of the habitat, have been proposed to affect welfare. While accredited facilities are required to meet high standards of care, habitat characteristics and management practices are not standardized across locations. Knowledge and subsequent application of these practices and habitat characteristics can enhance our understanding of factors associated with positive welfare. As part of a larger study of dolphin welfare titled "Towards understanding the welfare of cetaceans in zoos and aquariums" (colloquially called the Cetacean Welfare Study), survey data were collected from 86 bottlenose dolphins in 40 habitats at 38 facilities in seven countries. The major aims of this paper are to provide general descriptive information regarding dolphin management in accredited zoos and aquariums and to provide supplemental context to the other research published from the Cetacean Welfare Study data set. This paper provides a review of current habitat characteristics and management practices at those 38 accredited zoos and aquariums. These data enabled the identification and quantification of how cetacean management practices differed between participating facilities accredited by the Alliance for Marine Mammal Parks and Aquariums and the Association of Zoos & Aquariums. Variables were selected based on their potential association with welfare including the physical habitat, environmental enrichment, and training programs. The variables were also used for subsequent research in this collection of related papers to investigate important connections between potential indicators of welfare and habitat characteristics, environmental enrichment, and training programs.

Assessment of animal management and habitat characteristics associated with social behavior in bottlenose dolphins across zoological facilities.

Bottlenose dolphins are a behaviorally complex, social species that display a variety of social behaviors. Because of this, it is important for zoological facilities to strive to ensure animals display species-appropriate levels of social behavior. The current study is part of the multi-institutional study entitled "Towards understanding the welfare of cetaceans in zoos and aquariums" commonly referred to as the Cetacean Welfare Study. All participating facilities were accredited by the Alliance of Marine Mammal Parks and Aquariums and/or the Association of Zoos and Aquariums. Behavioral data were collected on 47 bottlenose dolphins representing two subspecies, Tursiops truncatus and Tursiops aduncus, at 25 facilities. The social behaviors of group related activity (group active) as well as interacting with conspecifics (interact with conspecific) were examined for their relationships to both animal management factors and habitat characteristics. The behavioral state of group active and the rate of interact with conspecific were both positively related to the frequency of receiving new forms of environmental enrichment. Both were inversely related to the random scheduling of environmental enrichment. Additional results suggested interact with conspecific was inversely related with daytime spatial experience and that males displayed group active more than females. Overall, the results suggested that animal management techniques such as the type and timing of enrichment may be more important to enhance social behavior than habitat characteristics or the size of the habitat. Information gained from this study can help facilities with bottlenose dolphins manage their enrichment programs in relation to social behaviors.

Relationships between animal management and habitat characteristics with two potential indicators of welfare for bottlenose dolphins under professional care.

Accredited zoos and aquariums continually strive to ensure high levels of animal welfare for the animals under their professional care. Best management practices include conducting research to better understand factors that lead to optimal welfare and then turning findings into practice. The current study is part of the larger Cetacean Welfare Study or more formally, "Towards understanding the welfare of cetaceans in zoos and aquariums." Facilities participating in the study were accredited by the Alliance of Marine Mammal Parks and Aquariums and/or the Association of Zoos and Aquariums. Animal management factors and habitat characteristics were examined in relation to two potential indicators of welfare for common (Tursiops truncatus) and Indo-Pacific (Tursiops aduncus) bottlenose dolphins. Specifically, we examined environmental enrichment, animal training, and habitat characteristics that were significantly related to behavioral diversity and route tracing, a form of stereotypic behavior. Behavior was recorded from 47 animals at 25 facilities around the world. Overall, the rate of route tracing behavior observed during the study was very low and few animal management factors or habitat characteristics were found to be related to this behavior. One factor, enrichment diversity, had a significant positive relationship with route tracing and an inverse relationship with behavioral diversity. This finding may be a product of a response mounted by animal care specialists to the behavior as opposed to a cause. Animals that engaged in this behavior were likely provided more diverse enrichment in attempts to ameliorate the stereotypic behavior. However, multiple factors were found to significantly relate to behavioral diversity, a potential positive indicator of welfare for bottlenose dolphins. Dolphins that were trained on a predictable schedule had higher behavioral diversity than those on a semi-predictable schedule. There was a positive significant relationship between behavioral diversity and the number of habitats to which an animal had access, and a significant inverse relationship with the maximum depth of the habitat. Finally, animals that were split into groups and reunited or rotated between subgroups had higher behavioral diversity than animals managed in the same group. Information gained from the current study suggested that animal management techniques may be more important in ensuring good welfare for bottlenose dolphins than focusing on habitat size.

Towards understanding the welfare of cetaceans in accredited zoos and aquariums.

Cetaceans are long-lived, social species that are valued as ambassadors inspiring the public to engage in conservation action. Under professional care, they are critical partners with the scientific community to understanding the biology, behavior, physiology, health, and welfare requirements of this taxonomic group. The Cetacean Welfare Study was a highly collaborative research effort among zoos and aquariums accredited by the Alliance for Marine Mammal Parks and Aquariums and/or the Association of Zoos & Aquariums that provided important empirical and comparative information on the care and management of cetaceans. The goal was to identify factors that were related to the welfare of bottlenose dolphins and to develop reference intervals and values for common and novel indicators of health and welfare for common bottlenose dolphins (Tursiops truncatus), Indo-Pacific bottlenose dolphins (Tursiops aduncus), beluga whales (Delphinapterus leucas), and Pacific white-sided dolphins (Lagenorhynchus obliquidens). Data were collected from cetaceans at 43 accredited zoos and aquariums in seven countries in 2018 and 2019. This overview presents a summary of findings from the initial research articles that resulted from the study titled "Towards understanding the welfare of cetaceans in zoos and aquariums." With multiple related objectives, animal-based metrics were used to advance frameworks of clinical care and target key conditions that were associated with good welfare of cetaceans in zoo and aquarium environments. As a result of this collaboration, species-specific reference intervals and values for blood variables and fecal hormone metabolites were developed and are freely available in an iOS application called ZooPhysioTrak. The results suggested that environmental enrichment programs and social management factors were more strongly related to behaviors likely indicative of positive welfare than habitat characteristics for common and Indo-Pacific bottlenose dolphins. These findings can be widely applied to optimize care and future science-based welfare practice.

Habitat characteristics and animal management factors associated with habitat use by bottlenose dolphins in zoological environments.

The way an animal uses its habitat can serve as an indicator of habitat appropriateness for the species and individuals. Bottlenose dolphins (Tursiops truncatus and Tursiops aduncus) in accredited zoos and aquariums experience a range of habitat types and management programs that provide opportunities for dolphins to engage in species-appropriate behaviors and potentially influence their individual and group welfare. Data in the present study were collected as part of a larger study titled "Towards understanding the welfare of cetaceans in zoos and aquariums" (colloquially called the Cetacean Welfare Study). Non-invasive bio-logging devices (Movement Tags) recorded the diving behavior and vertical habitat movements of 60 bottlenose dolphins at 31 zoos and aquariums that were accredited by the Alliance for Marine Mammal Parks and Aquariums and/or the Association of Zoos & Aquariums. Bottlenose dolphins wore a Movement Tag one day per week for two five-week data collection periods. Demographic variables, environmental enrichment programs, training programs, and habitat characteristics were associated with habitat usage. Longer dive durations and use of the bottom third of the habitat were associated with higher enrichment program index values. Dolphins receiving new enrichment on a monthly/weekly schedule also used the bottom third of the habitat more often than those receiving new enrichment on a yearly/year+ schedule. Dolphins that were managed in a group that was split into smaller subgroups during the day and were reunited into one group at night spent less time in the top third of the habitat than those who remained in a single group with consistent members at all times. Dolphins that were managed as subgroups with rotating members but were never united as one group spent less time in the bottom third of the habitat than those who remained in a single group with consistent members at all times. Taken together, the results suggested that management practices, such as enrichment and training programs, played a greater role in how dolphins interacted with their environment relative to the physical characteristics of the habitat.

Behavioral diversity as a potential positive indicator of animal welfare in bottlenose dolphins.

Accredited zoological facilities are committed to fully understanding the behavioral, mental, and physical needs of each species to continuously improve the welfare of the animals under their professional care and detect when welfare has diminished. In order to accomplish this goal, internally consistent and externally valid indicators of animal welfare are necessary to advance our understanding of the current welfare status of individual animals. Historically, efforts have focused on monitoring visible or observable signs of poor health or problem behavior, but lack of signs or problems does not necessarily demonstrate that an individual animal is thriving. The current study examined fecal hormone metabolite levels and behavior for two species of bottlenose dolphins (Tursiops truncatus and Tursiops aduncus) from 25 different accredited zoological facilities. At the time of the study, all facilities were accredited by the Alliance of Marine Mammal Parks and Aquariums and/or the Association of Zoos and Aquariums. This was part of the multi-institutional study 'Towards understanding of the welfare of cetaceans in zoos and aquariums" commonly referred to as the Cetacean Welfare Study. Behavioral diversity was calculated using the Shannon Diversity Index on species-appropriate behavioral events. Behavioral diversity was compared to the fecal metabolites of cortisol, aldosterone, and the ratio of cortisol to dehydroepiandrosterone (DHEA) as well as the stereotypic behavior of route tracing. Similar to previous studies on other species, there was a significant inverse relationship between behavioral diversity and both fecal cortisol metabolites and route tracing. Additionally, a significant inverse relationship also exists between behavioral diversity and the ratio of fecal cortisol to DHEA metabolites. Behavioral diversity and fecal aldosterone metabolites were not associated. Additional research is still needed to validate behavioral diversity as an indicator of positive animal welfare for bottlenose dolphins and across species. However, based on current results, facilities could utilize behavioral diversity combined with other measures of welfare to more comprehensively evaluate the welfare of bottlenose dolphins.