This article describes and discusses a set of exercises in silent zoo visiting conducted at two zoos in the Southwest of the UK in 2019. The visits were part of a wider project on the auditory culture of zoos, institutions which have tended to be analysed by reference to their concern with the visual display of captive other-than-human animals (henceforth animals). Linking ideas on silence that have emerged from work in anthropology and sound studies respectively, the article explores the notion that among many other affordances, silence can generate opportunities for reflection and the consideration of alternatives, in this case in relation to a prevailing zoo visiting culture often characterised by loud anthropogenic sound. Examining extracts and observations from discussions among participants, the article illustrates how this novel form of visiting fostered a variety of types and qualities of experience. It also provided an impetus for participants to notice and focus on specific animals and aspects of animal lives they might not otherwise have considered. Such meditation could foster feelings of connection to particular zoo animals, while also opening up possibilities for more detached contemplation. The silent visits led participants to produce critical perspectives on conventional zoo visiting, and to imagine future possibilities for the auditory culture of zoos. The article shows that silence can serve as a productive methodological tool in the exploration of human-animal relations, and demonstrates how sound is integral to the formation of those relations within, and by implication beyond, the zoo.

Flamingos are colonial species commonly kept in zoos, well known for their bright plumage and elaborate courtship displays. This project aimed to determine the differences in flock position and association preferences of juvenile Greater Flamingos (Phoenicopterus roseus) and Caribbean Flamingos (P. ruber) housed in the same zoological collection. Little research has been conducted on the association preferences of juvenile flamingos, especially in captive flocks, and therefore this study collected data using photographs taken throughout 2014 and 2015 to further understand association patterns. Data were collected on the age category of each juvenile flamingo observed, the age of their nearest neighbour and their position within the flock, and the location within an enclosure zone at different times of the day. The results showed that Greater Flamingo juveniles mainly associated with individuals of their own age and were most likely positioned at the periphery of their flock significantly more of the time until approximately 24 months of age. Sub-adult Greater Flamingos spent significantly more time associating with adult flamingos at the centre of the flock. In contrast, data collected on Caribbean Flamingos indicated that juveniles did not segregate themselves from the adults as distinctively. Birds aged 13–24 months were observed significantly more at the centre of the flock and had more associations with adult flamingos, in a similar manner to that observed in Greater Flamingos. Due to population management needs, juvenile Caribbean Flamingos were removed from the flock at the start of 2015 and this may have influenced the association and location preferences of the remaining young flamingos. In conclusion, these results indicated that captive juvenile flamingos were often seen away from adult birds and that sub-adult flamingos returned to the heart of their natal flock to associate significantly more with other adult individuals, potentially preparing for mate selection and breeding. Captive enclosure should therefore be spacious enough to enable young flamingos to remove themselves from adult birds so that behavioural development can be unaffected by artificially high rates of aggression.

AbstractEvidence-based approaches are key to advancing all areas of zoo and aquarium practice. Output from empirical study must be disseminated to those within the industry so that results can support changes to husbandry and management for individual species. Information on enrichment techniques is published in a range of sources, including papers in the peer-reviewed and ‘grey literature’ (ie professional but non-reviewed publications). To investigate how evidence is implemented into enrichment practices, we sampled all enrichment studies identified in one online repository of peer-reviewed papers and two grey literature publications across an eleven-year period. We recorded whether the enrichment was supported with biological evidence (whether it was developed using published enrichment-focused research for that species and/or with the species’ ecology and behaviour information) alongside analysis of the type of enrichment used and the chosen study species. Enrichment articles were more likely to be supported by biological evidence in peer-reviewed than grey literature. Taxonomic differences in the use of evidence were noted; for example, enrichment provided to carnivores and parrots was more likely to be supported with biological evidence compared to that used for penguins. of the five enrichment types, nutritional enrichment was most often based on biological evidence. Multi-category and physical enrichment types were more common across all literature sources whereas social enrichment was less common, suggesting barriers to implementation of all enrichment types in zoological facilities. Our research suggests that zoo and aquarium professionals are considering species-specific welfare needs by ensuring that enrichment protocols are supported by biological evidence. However, opportunities to diversify the enrichment types being offered and species being researched are identified.

AbstractThe implementation of environmental enrichment (EE) can be effective in promoting zoo animal welfare by enhancing the performance of natural or species‐typical behaviors. Research on the effects of EE is biased towards larger mammalian species, with less charismatic species being overlooked. Armadillos are one such overlooked example. A captive environment that results in inactivity, obesity, and associated poor health can negatively affect armadillo well‐being. The aim of this study was to evaluate how the implementation of four physical object‐based EEs could positively affect the behaviors of three armadillo species, housed in four similar exhibits. Behavioral data were collected both before (baseline) and during (treatment) EE periods, alongside of visitor number and environmental temperatures. The EE comprised of a plastic ball or a cardboard tube or a cardboard box, or a scatter‐feed, and these were rotated each week of study until each exhibit had received them in turn. Despite the presence of different EE types, activity remained low throughout the study. However, results suggest that the plastic ball and cardboard box increased exploratory behaviors in the armadillos, but no overall increase in activity was noted during the scatter feed. Visitor presence had no effect on armadillo activity, and armadillos showed reduced activity with increasing environmental temperature. Overall, the use of physical object‐based EE promoted beneficial natural behaviors in zoo‐housed armadillos, but environmental conditions (i.e. temperature) also impacted armadillo activity, suggesting a complicated relationship between an enclosure's environmental variable and any behavioral husbandry measures.

Foot lesions are a highly prevalent phenomenon among zoo-housed flamingos, with up to 99.8% of birds affected. These lesions are a recognized welfare concern, increasing the likelihood of bacterial infections, and even septicemia. Although several risk factors have been linked to foot lesions in flamingos (including age, climate, and substrate), there have been few studies looking at changes in foot lesions over time. This study tracked changes in foot lesions for an individual flock of Chilean Flamingos (97 birds) at Dublin Zoo, Ireland, over an 18-month period in response to a mandatory indoor housing order imposed by the Irish Government as a seasonal precautionary measure to prevent the spread of avian influenza. Using a pre-defined scoring system for four common types of foot lesions (hyperkeratosis, fissures, nodular lesions, and papillomatous growths), we show that providing unrestricted access to outdoor habitats and natural substrates (both terrestrial and aquatic) can improve the health and wellbeing of zoo-housed flamingos. This longitudinal study highlights the importance of regular foot health monitoring in flamingos, and the importance of natural aquatic substrates when managing flamingos. As many zoo-housed birds have been spending more time indoors on artificial substrates over recent years due to avian influenza housing orders, it is critical that we assess the impact of such changes in management and habitat access on bird health and welfare.

Good mental and physical health go hand-in-hand when identifying factors that lead people to experience a better overall quality of life. A growing disconnect to the natural world is worsening the mental health of individuals in many societies. Numerous scientific publications have evidenced that being in nature and access to green and blue spaces positively impact upon humans’ physical and mental health. For many people, particularly those living in more urbanized areas, managed natural spaces and borrowed landscapes, such as those found in public parks, wildlife reserves and zoological gardens give the only opportunities for wider engagement with nature. Many zoos are designated green spaces and therefore human visitors can engage with native fauna and flora as well as exotic wild animals. This article reviews the UK Government’s “The Five Ways to Wellbeing” concept, applied to zoos and aquariums and thus suggests how zoos and aquariums can use this framework to promote positive nature-connectivity experiences for their visitors and promote good wellbeing. The Five Ways to Wellbeing are Connect, be active, Take notice, Keep learning, and Give. We illustrate how zoos and aquariums could model their approaches to educational and engagement roles, as well as design initiatives to reach out to local communities via the Five Ways to Wellbeing concept. We show that many of the positive programs and works conducted by zoos and aquariums lend themselves to further engagement with the Five Ways to Wellbeing structure. By taking such a structured approach in the design, implementation and evaluation of their activities, zoos can expand their abilities in connecting humans with nature and further add value to their living collections of animals and plants. By including Wellbeing as a defined aim of the modern zoo, it will be clear to all of those involved in their work, visitors, workers, stakeholders, that zoos are working to promote, protect and preserve positive wellbeing outputs for humans and animals alike.

AbstractAs global wildlife populations continue to decline, the health and sustainability of ex situ populations in zoos and aquariums have become increasingly important. However, the majority of managed ex situ populations are not meeting sustainability criteria and are not viable in the long term. Historically, ex situ flamingo (Phoenicopteriformes) populations have shown low rates of reproductive success and improvements are needed for long‐term viability. Both flock size and environmental suitability have previously been shown to be important determinants of ex situ flamingo reproductive success in a limited number of sites in some species. Here we combined current and historic globally shared zoological records for four of the six extant species of flamingo (Phoeniconaias minor, Phoenicopterus chilensis, Phoenicopterus roseus, and Phoenicopterus ruber) to analyze how flock size, structure, and climatic variables have influenced reproductive success in ex situ flamingo populations at 540 zoological institutions from 1990 to 2019. Flock size had a strong nonlinear relationship with reproductive success for all species, with flock sizes of 41–100 birds necessary to achieve ca. 50% probability of reproduction. Additionally, an even sex ratio and the introduction of new individuals to a flock both increased ex situ reproductive success in some cases, while climatic variables played a limited role. We demonstrate the conservation management potential from globally shared zoological data and provide species‐specific management recommendations to increase the reproductive success of global ex situ flamingo populations: minimum flock sizes should be increased, and we encourage greater collaboration between individual institutions and regional associations in exchanging birds between flocks.

Measurement of zoo animal welfare states enables improvement to husbandry and management to be evidence-based and implemented according to species’ needs. Theoretical welfare concepts are often discussed, and whilst it is helpful to ensure wide consensus across all stakeholders on what welfare comprises, practical application of such evidence-based information is as equally important. All species housed in zoos will have specific needs that must be met by their housing, husbandry, and care to enable them to thrive. Therefore, this paper examined how to identify key animal care needs and an animal’s responses to them to form a basis for species-specific welfare assessment approaches. There are examples of familiar-to-the-zoo species that still pose challenges regarding delivery of optimal husbandry and management. As such, the identification and evaluation of core concepts of the biology, behaviour, “needs and wants” of these species is required to support validation and refinement of physical, behavioural, and psychological welfare measures. This article evaluated the use of evidence to build capacity in welfare measurement for a familiar zoo-housed species, the giraffe (Giraffa camelopardalis, Linnaeus 1758), by outlining seven key needs and requirements that must be provided in the zoo (“giraffe W-E-L-F-A-R-E” = Warmth, Enrichment, Leaves, Feeding, Alfalfa, Rumination, Exercise). Provision of these inputs, and opportunities for all giraffes in a herd to engage with them, provides the foundation for further welfare assessment to be implemented. Specifically, the validation and measurement of mental states that are more likely to be positive if key behavioural and ecological needs have already been met. This paper advocated for this evidence-based approach to “welfare-focussed husbandry”, with distillation of key information that supports species-relevant care, to be developed for other zoo-housed species as support for their welfare assessment protocols. Such welfare-focussed husbandry is layered on top of the basic animal care requirements of the species in the zoo to ensure all individuals have the best opportunity to attain positive welfare states. In this way, and once validated, foundational welfare assessment can be easily completed by busy animal care staff, capacity is built into zoo operations as all stakeholders are aware of exactly what each species needs, and deeper dive welfare assessment (especially concerning animal mental states) can be targeted more effectively. Further evolution of these seven steps for the giraffe is suggested and extrapolation of this approach, to aid identification of key welfare indicators across all zoo-housed species, is encouraged.

AbstractAnimal personalities manifest as consistent individual differences in the performance of specific behavioural expressions. Personality research has implications for zoo animal welfare, as it can further our understanding of how captive individuals may differ in their resource use and provide insight into improving individual and group social health. For group living species, personality may enable assortment based on similar behaviour and influence an individual’s interactions with conspecifics (e.g. social support). This research aimed to document how personality traits (aggressive, exploratory, submissive) influenced the social network structure of highly social animals in a captive environment. Data were collected from separate flocks of captive Caribbean (Phoenicopterus ruber) and Chilean flamingos (Phoenicopterus chilensis) to identify relationships between birds and examine opportunities for social support. The flocks associated non-randomly, and in both cases, personality was a substantial predictor of network structure. Personality also predicted key elements of Caribbean flamingo social role (degree, betweenness and average association strength) conflict outcome, and propensity to provide social support, however these patterns were not replicated within the Chilean flamingo network. While both species appear to assort by personality, the broader relationship between personality and social role may vary depending on species and context.

AbstractEnvironmental enrichment (EE) is commonly provided to animals managed under human care, being beneficial to behavioral diversity and improving animal welfare. Use of EE appears to be particularly beneficial to individual wild animals spending a short period of time in captivity, for example, as part of conservation or rehabilitation programs. This paper documents a case study on the application and relevance of EE for a group of captive mute swans housed in a rescue center. Observational data were analyzed for two groups of juvenile swans that were provided with a physical EE device to increase time spent foraging. Periods of no EE were observed and compared to data from when birds were provided with EE. Results show that EE promoted foraging time and helped to reduce long periods of inactivity in captive birds. EE helped to reduce occurrence of captive‐focused (i.e. abnormal behaviors) although these was already seen at very low rates. Inactivity as a measure of welfare in captive swans specifically (and waterbirds generally) should be further investigated to understand potential impacts on bird health. Our research shows the benefits of simple and easy‐to‐use EE devices on captive animal behavior and how use of EE for individuals spending a short amount of time in captivity (e.g. within a rescue center) could ensure diversity of behavior patterns and promote the performance of adaptive behaviors upon release to the wild.

The Serengeti-Maasai Mara ecosystem is a dynamic environment that is home to some of the last megafauna on earth, the world’s largest mammalian migration, and a diversity of human subsistence strategies. This research focuses on the spread of the disease malignant catarrhal fever from wildebeest calves in the northern part of the ecosystem to cattle belonging to Maasai pastoralists and how subsequent loss of cattle and avoidance of diseased areas affects Maasai livelihoods, attitudes toward wildebeest and conservation, and the behavior of wildebeest in the area. Understanding the attitudes local Maasai hold toward wildebeest and conservation, and the consequences of those attitudes on conservation success and wildebeest behavior, is critical to successful conservation of the wildebeest and the ecosystem that depends on them.

To investigate the attitudes and behaviors of Maasai pastoralists toward wildebeest and conservation, an online survey of 114 Kenyan Maasai people was conducted using social media. In order to inform the interpretation of attitudes on conservation, an email survey of 16 conservation practitioners who had worked or were currently working in Africa was conducted. In order to determine possible differences in wildebeest behavior in Maasai versus non-Maasai inhabited areas, behavioral data (collected by the Snapshot Serengeti Project and analysed by citizen scientists) in Enonkishu Conservancy in Kenya (a Maasai area) and Serengeti National Park in Tanzania (where Maasai are largely excluded) were compared.

The results of the survey of Maasai people indicated high levels of disturbance and livelihood loss due to the presence of wildebeest infected with malignant catarrhal fever. Respondents reported broad dislike and avoidance of wildebeest that strained their livelihoods and a strong desire for a malignant catarrhal fever vaccine. The survey of conservation practitioners indicated a great desire to work with local people and a broad belief that respondents were currently doing so. However, the conservation practitioner survey indicated involvement of local people as mainly peripheral members of conservation research—as drivers or guides. The Maasai survey indicated a desire among Maasai respondents to be involved on a deeper level with conservation (e.g. as educators and members of the research team). The wildebeest behavioral data showed wildebeest in Enonkishu Conservancy had significantly lower rates of standing, resting, eating, and interacting and significantly higher rates of moving. Though further study is needed to determine the exact cause, it does indicate a difference in behavior between the two sites.

Though this study was limited in the sample sizes of the two surveys and in the multiple differences between the two sites used for the wildebeest behavioral analysis, it has helped expound on the conservation encounter between wildebeest, Maasai pastoralists, and conservation efforts in Kenya. These results suggest that more cooperation between local Maasai and conservation personnel is needed for more effective conservation, and that a vaccine for malignant catarrhal fever could help support Maasai livelihoods. This study has also indicated that Maasai presence could be affecting wildebeest behavior. This could be critical in investigating the degree that wildebeest may adapt to changing conditions, such as human encroachment and climate change.

Modern zoos and aquariums have evolved greatly since the end of the Second World War, to become centres of conservation excellence and scientific institutions for the study of animal behaviour, ecology, husbandry management. Whilst the impact of zoos and aquariums to biodiversity conservation, population management and advancement of species care is well documented, their positive impacts on society (including the communities that they are located within) is less well known. The four established aims of the modern zoo—Conservation, Education, Recreation (Engagement) and Research—provide a strong foundation for wider review and critique of the societal value of zoos and aquariums. This article synthesises what such value may be, and how it could be measured, from a systematic review of the literature pertaining to each of these four established aims. It also recommends areas of future scientific inquiry to further study the wider impact of zoos on their local communities and on human populations and behaviour more generally. Including Wellbeing as a fifth aim of the modern zoo would also cement their wider societal value. Further scrutiny of the societal value of the modern zoo enhances the value of the zoo’s living collection and the green spaces that it manages to provide accessibility to biodiversity and nature-centric education essential to long-term, planetary friendly human behaviour change.

Mixed species exhibits in zoos are used to create larger, more stimulating environments to support naturalistic interactions between species. In the wild, mixed species groups are observed as having lower rates of vigilance, presumably due to reduced predation risk through ’detection’ and ’dilution’ effects. This effect appears to be highly variable depending on factors such as food availability or degree of threat. This study aimed to collect data on mixed-species associations and consequent vigilance rates in the wild, collecting equivalent data from a large mixed-species zoo enclosure to compare the findings between free-ranging and captive populations. The study additionally investigated whether large mixed-species enclosures support natural associations and behaviours, by comparing the behaviour of captive animals with wild counterparts. The study used livestream video feeds from 10 national parks in South Africa and Kenya to observe free-ranging species, and a camera at the San Diego Zoo Safari Park’s mixed species African exhibit. Scan and continuous sampling protocols were used simultaneously to record behavioural states as well as the rate of scanning (vigilance) events. GLMMs were run to test whether vigilance of a focal species varied according to the number of animals present, the density of animals in the group, and the diversity of species. In the wild, vigilance decreased with increasing number of animals in the surroundings but in captivity the group size had no impact. The results suggest that in the wild, these species benefit from increased perceived safety in larger groups, regardless of the species making up that group. No effect was noted in the zoo because of a reduced need for animals to show heightened vigilance to the same degree as in the wild. Similarities were observed in associations between species/mixed species group compositions, and in behaviour budgets. These findings provide a preliminary evaluation of how the impact of mixed species groupings may translate from the wild to the zoo, based on the associations and behaviour across a variety of African ungulates.

Zoo management techniques for captive birds, such as flight restraint and enclosure type, may affect behavioral performance and are consequently worthy of investigation. Flamingos are amongst the most popular of zoo-housed birds and, as such, research into their captive management and associated behavioral responses are widely applicable to many thousands of individuals. As a highly social species, understanding social bonds and behavior of the individual bird and the flock overall can help inform decisions that support husbandry and population management. In this project, 41 greater flamingos at Bristol Zoo Gardens were observed for 49 days across spring and summer 2013 to assess the following: (i) social associations within the flock, (ii) overall activity patterns, and (iii) distribution of time within specific enclosure zones for both full-winged and flight-restrained birds living in the same enclosure. Results showed that pinioning interacted with age in regard to flamingo time-activity patterns, but wing condition did not significantly influence association patterns, performance of social interactions, or performance of breeding behavior. Social network analysis revealed that associations were nonrandom and flamingos, of either wing condition, displayed different roles within the network. Birds of similar age formed the strongest bonds. Enclosure usage was not even, suggesting that the flamingos favored specific areas of the enclosure during the observation period. This study showed that wing condition does not affect flamingo behavior, social bonds, or space use, and that age and sex have more of an overall influence on what flamingos do, and with whom they chose to do it. Further research should extend this study into other, larger captive flocks to further refine behavioral measures of welfare for these popular zoo birds.

Individual animals engage in many behaviours which are mutually exclusive, and so where individuals increase the duration of time spent on one type of behavioural activity, this must be offset by a corresponding decrease in at least one other type of behaviour. To understand the variation observed in animal behaviour, researchers need to know how individuals trade-off these mutually-exclusive behaviours within their time-activity budget. In this study, we used remotely collected behavioural observations made from a live-streaming webcam to investigate trade-offs in the behaviour of two bird species, the mute swan (Cygnus olor) and whooper swan (Cygnus cygnus). For both species, we tested for correlations in the duration of time spent on key mutually exclusive behaviours: aggression, foraging, maintenance, and resting. We detected a negative association between aggression and resting behaviours in both species, indicating that increased aggression is achieved at the expense of resting behaviour. In contrast, there was no apparent trade-off between aggression and foraging, aggression and maintenance, or maintenance and resting. Foraging and resting behaviours were negatively correlated in both species, highlighting a trade-off between these distinct modes of behaviour. A trade-off between foraging and maintenance behaviours was detected for the sedentary mute swans, but not the migratory whooper swans. Our findings show how birds can trade-off their time investments in mutually exclusive behaviours within their time-activity budgets. Moreover, our study demonstrates how remotely-collected data can be used to investigate fundamental questions in behavioural research.

Data on zoo enclosure usage provide meaningful evaluation of husbandry and welfare but for social species, group-level data may not capture individual occupancy preferences. Determining zone occupancy using group data may give an inaccurate assessment of enclosure suitability for each individual. We compared three formulae (traditional and modified Spread of Participation Index and Electivity Index) to determine how estimations of space usage compare between individuals and their group overall. Two flamingo flocks at two zoos were sampled with enclosures separated into discrete zones. Counts of where each bird in the flock was located at each minute of sampling were compared against the number of minutes that randomly selected individual flamingos spent in each zone. Overall, there was little difference in preferred zone occupancy when flock data were compared to individual data. Group data suggested that flamingos were more consistent in their enclosure usage, with individual data showing wider overall usage of enclosures. Individual Electivity Index (EI) values suggested zone underuse whereas group EI suggested zone overuse. As a proxy for welfare measurement, we recommend both group-level and individual monitoring of space use to provide a complete picture of how individuals within a social group occupy their space and choose where to be.

Many species of birds are housed in zoos globally and are some of the most popular of animals kept under human care. Careful observations of how species live and behave in their natural habitats can provide us with important knowledge about their needs, adaptations, and internal states, allowing identification of those behaviours that are most important to the individual’s physical health and wellbeing. For this study, Mandarin Ducks (Aix galericulata) were chosen as a study species because, like many species of waterfowl, they are widely kept in both private institutions and zoos, yet little research has been conducted on their core needs in captivity. A free-living population of naturalised Mandarin Ducks living in Richmond Park was used for this research. Data on state behaviours (resting, swimming, foraging, perching, preening, and vigilance) were collected five days a week (08:00–18:00) from the 26 March to 26 May 2021. Secondly, temporal, seasonal, environmental, and animal-centric factors (e.g. Sex) were recorded to assess any impact on the Mandarin’s time-activity budget. Lastly, a comparison between free-living anmd captive activity was conducted (via the literature) to evaluate whether captive behaviours differ to how they are expressed in the wild. Results showed that free-living Mandarins predominantly rested (19.88% ± 28.97), swam (19.57% ± 19.43) and foraged (19.47% ± 25.82), with variations in activity related to factors such as vegetation cover and pond size. Results also showed differences between the time-budgets of free-living and captive Mandarins, suggesting that captive birds may not always have the opportunity to express species-typical behaviours. This research indicated that study of natural behaviours performed in the wild may help to evaluate “normal” behaviour patterns of zoo-housed individuals and provide evidence for environmental and husbandry alterations that can promote good welfare. However, any potential impact on the activity patterns of free-living species due to human interactions should be considered when assessing deviations between the behaviour of wild and captive individuals.

Zoos and aquariums have an enormous global reach and hence an ability to craft meaningful conservation action for threatened species, implement educational strategies to encourage human engagement, development and behavior change, and conduct scientific research to enhance the husbandry, roles and impacts of the living collection. The recreational role of the zoo is also vast- people enjoy visiting the zoo and this is often a shared experience amongst family and friends. Evaluating how the zoo influences this “captive audience” and extending its reach to include a compassionate approach to animals and people can further enhance the mission, value, and relevance of their work. The modern zoo’s current aims—Conservation, Education, Research and Recreation—provide useful foundations for the activities that zoos conduct at a local and national level. However, to improve sustainability of their actions and outcomes, we feel that Wellbeing should become the fifth aim of the modern zoo for the future- both from an animal perspective (ensuring that populations are managed according to prevailing behavioral needs) and from a human perspective (enhancing access to nature, promoting planetary friendly behavior changes and ways of living, and advancing the wellbeing of the zoo’s workforce). This paper provides discussion and review of how Wellbeing is already a substantial part of what zoos work on as well as posing the idea of altering the Recreation aim of the zoo to one of Engagement, which potentially is more measurable and therefore can allow zoo researchers and managers further options for the collection of evidence on the local and global reach of their zoo’s aims. Education, Engagement, Conservation, Research and Wellbeing provide a more complete picture of the roles of the modern zoo for the animals (bothin situandex situ), human visitors and workforce, and to society more widely.

Visitors can influence the behaviour of zoo animals through their auditory and visual presence, with mixed findings of negative, neutral, and positive effects on welfare. This study opportunistically utilised the UK-wide COVID-19 period of zoo closure to investigate the activity and enclosure usage of Greater (Phoenicopterus roseus) and Chilean (P. chilensis) Flamingos housed at two zoos. Flamingo behaviour at both sites was observed during the last week of a three-month closure period and the immediate reopening of the zoos. Photographic data were collected at three timepoints during each observation day. Negative binomial GLMMs compared the behaviour observed during zoo closure to the behaviour observed during zoo reopening, whilst accounting for climatic variables and time of day. Spearman’s correlation identified relationships between behaviour with the number of visitors and weather. Greater Flamingos were not influenced by the reintroduction of visitors to the zoo setting. Chilean Flamingos showed an increase in inactivity and decrease in movement and feeding when the zoo reopened. These possible behavioural responses are better explained by the influence of temperature on the behaviour of Chilean Flamingos and by the correlation between temperature and visitor number, rather than a direct consequence of visitor presence. This research details the multifactorial nature of any potential anthropogenic effects on zoo animal behaviour and highlights the importance of considering environmental variables alongside the measurement of visitor presence or absence.

Flamingos (Phoenicopteriformes) are obligate colonial species that nest in large colonies, with monogamous pairs rearing a single chick following a synchronised group courtship display. Within this relatively simplistic behavioural description, deviations from these social and reproductive norms are apparent. Same sex pairings, multi-bird relationships and extra pair copulations are documented in the literature. Flamingos display highly sexually selected characteristics of plumage colour, carotenoid accumulation and diversity of display movements that underpin mate choice decisions. The brightest birds in best body condition are more successful at breeding. Therefore, documented mate guarding of female birds by male partners, is a relevant response to maximise investment in a pair bond. Limited information that describes the action of forced copulation by the male flamingo and the response of the female bird is available in the literature. This paper describes an occurrence of an attempted forced copulation by an older male Greater Flamingo to a younger female bird. Such behaviour may be an artefact of the captive environment, and limited mate choice when compared to the sizes of wild flocks, or it could be regularly apparent in the wild and therefore worthy of more scrutiny and empirical study.

The study of animal behaviour is important for the development of husbandry and management practices for zoo-housed species. Yet, data are typically only collected during daylight hours, aligning with human work schedules rather than animal activity patterns. To remedy this, 24 h data collection is needed. This study investigated the behaviour of a captive flock of lesser flamingos to understand temporal changes in their time-activity patterns. Two remote camera traps were placed around the birds’ outdoor enclosure and one within the indoor house. Counts of birds visible within specific enclosure zones were recorded from photographic data. Behaviour was defined as active or inactive, and modified Spread of Participation Index (SPI) was used to calculate enclosure zone occupancy. Results indicated that lesser flamingos are active overnight, and to a similar amount as in the daytime. Proportions of birds observed as active were significantly higher at later times of the day (i.e. dusk) when compared to the number of active birds in the morning. Enclosure usage was diverse and indoor and outdoor zones could be used by different numbers of birds at different times of the day. Variation in enclosure usage may indicate the changing needs of the flamingos when housed indoors overnight and when they have night-time access to an outdoor enclosure. This research has identified the need for further research into the nocturnal behaviour and space use of lesser flamingos and suggests the need for 24 h research in captive birds, and other zoo-held species, especially when species are locked indoors or face behavioural restriction overnight due to biosecurity measures surrounding zoonoses outbreaks, e.g. Avian Influenza.

The fossa is a specialized Malagasy carnivore housed in ex situ facilities since the late 19th century. Moderate breeding success has occurred since the 1970s, and welfare issues (notably stereotypic pacing behaviour) are commonly documented. To understand challenges relating to fossa housing and husbandry (H&H) across global facilities and to identify areas of good practice that dovetail with available husbandry standards, a survey was distributed to ZIMS-registered zoos in 2017. Results showed that outdoor housing area and volume varied greatly across facilities, the majority of fossa expressed unnatural behaviours, with pacing behaviour the most frequently observed. All fossa received enrichment, and most had public access restricted to one or two sides of the enclosure. The majority of fossa were locked in/out as part of their daily management and forty-one percent of the fossa surveyed as breeding individuals bred at the zoo. Dense cover within an enclosure, restricted public viewing areas, a variable feeding schedule and limited view of another species from the fossa exhibit appear to reduce the risk of unnatural behavior being performed. The achievement of best practice fossa husbandry may be a challenge due to its specialized ecology, the limited wild information guiding captive care, and the range of housing dimensions and exhibit features provided by zoos that makes identification of standardized practices difficult. We recommended that holders evaluate how and when enrichment is provided and assess what they are providing for environmental complexity as well as consider how the public views their fossa.

Amphibians are an understudied group in the zoo-focussed literature. Whilst commonly housed in specialist exhibits and of real conservation value due to the global extinction crisis, amphibian welfare is not often investigated empirically in zoo settings. The limited research that is available suggests that enclosure design (structure, planting and naturalistic theming) has a positive impact on the time that amphibians will be on show to visitors. However, the categorisation of any “visitor effect” (i.e. influences of visitor presence on amphibian activity and time on display) is hard to find. The COVID-19 pandemic forced the closure of zoological organisations in the UK for several months from March 2020, with gradual re-openings from the summer into autumn and winter. This event provided a unique opportunity to study the effect of the lack of visitors, the presence of essential zoo staff only, the wider return of organisational staff, and then the return of visitors over a prolonged period. This project at WWT Slimbridge Wetlands Centre assessed the number of individuals of six species of amphibian—common toad (Bufo bufo), common frog (Rana temporaria), smooth newt (Lissotriton vulgaris), pool frog (Pelophylax lessonae), golden mantella (Mantella aurantiaca) and golden poison dart frog (Phyllobates terribilis)—visible to observers under different conditions. All amphibians were housed in a purpose-built indoor exhibit of individual enclosures and were recorded when visible (as a proportion of the total population of the enclosure) during closure, the return of extra centre staff and visitor periods. The results showed species-specific differences in visibility, with some species of amphibian being more likely to be on view when the presence of people at their enclosure was less likely or in smaller numbers. Such differences are likely related to the specific camouflage or anti-predation tactics in these focal species. Further study to quantify amphibian sensitivity to, and perception of, environmental change caused by public presence (e.g. light levels and sound) would be useful welfare-themed research extensions. Our results can help inform husbandry, collection planning and amphibian enclosure design to reduce any noticeable visitor effects, and provide a useful benchmark for further, more complex, welfare assessment measures.

Behavioural research in zoos is commonplace and is used in the diagnosis and treatment of potential husbandry and management challenges. Robust methods that allow valid data collection and analysis constitute an evidence-based approach to animal care. Understanding behaviour is essential to improving animal management, and behavioural research is therefore popular, with a wide choice of behavioural methodologies and theories available. This review outlines ten methodological approaches, concepts or theories essential to zoo science that are based around behavioural observation. This list is not exhaustive but aims to define and describe key areas of consideration when planning and implementing a zoo-based behavioural project. We discuss the application of well-established methods (the construction of ethograms, use of time–activity patterns and measurement of space/enclosure use) as well as evaluating newer or less-widely applied analytical techniques, such as behavioural diversity indices, social networks analysis and Qualitative Behavioural Assessment. We also consider the importance of fundamental research methods, the application of pure science to understand and interpret zoo animal behaviour (with a review of a Tinbergian approach) and consideration of meta-analyses. The integration of observational techniques into experiments that aim to identify the cause and effect of behavioural performance is then explored, and we examine the assimilation of behavioural methods used in studies of environmental enrichment. By systematically studying animal behaviour, we can attempt to understand the welfare of individual animals in captivity, and here we present an example of our reviewed approaches to this area of zoo science. Combining multiple methodologies can lead to a greater understanding of behaviour and welfare, creating robust research, progressing husbandry and advancing conservation strategies. Collaborations between zoological collections and academic researchers (e.g. in Higher Education Institutions) can further refine and enhance the validity of research and husbandry practice alike.

YouTube is the dominant online video-sharing platform and offers zoos an opportunity to engage a vast audience with conservation content. As there is limited research evaluating how zoos currently utilize YouTube, we cataloged and evaluated the content of 20 zoological organizations (focusing on the top 1000 most viewed videos from each channel) from 2006 to 2019. Separately, educational content in a subsample of the most viewed and recently produced videos from three zoo-YouTube channels was catalogued and evaluated for the period 2016 to 2019. We found that channels are becoming more focused on producing entertainment, rather than education-based content and not reflecting the taxonomic diversity of their animals. Videos containing conservation content may not be the most popular with the YouTube audience (making up only 3% of the most viewed videos), but there is evidence that outputs are becoming more conservation orientated, with more conservation-focused videos being produced over time. Zoos may be engaging with large audiences via YouTube, but should carefully consider how they represent different taxonomic classes and how they can create educational material in interesting and accessible forms.

Behaviour signals the internal states that relate to an individual’s welfare and its development is influenced by the early social environment that an animal experiences. Husbandry practices can alter this early social environment, for example different rearing conditions (e.g. foster rearing by a surrogate parent of another species). Widespread implementation of altered rearing can lack empirical support and non-parent-reared animals may experience poorer welfare resulting from maternal deprivation. An opportunity presented itself to measure the effect of foster-rearing on Chilean flamingo behaviour and social preferences at WWT Slimbridge Wetland Centre and compare findings to parent-reared conspecifics in the same time period. Data were collected from April to July 2019 at three timepoints during each observation day. Binomial generalized linear mixed models were used to assess the relationship between focal chicks’ rearing background with behaviour, zone usage, and flock position whilst accounting for climatic factors and visitor numbers. The development of social preferences was assessed using social network analysis. Our results showed limited impacts on flamingo behavioural development due to foster rearing. Foster-reared chicks spent less time feeding, were more likely to occupy the nesting area of the enclosure, and had fewer significant preferred associations than parent-reared chicks, but preferred social bonds were as equally strong and durable for both foster-reared and parent-reared chicks. Our results have important welfare implications for the use of foster-rearing in captive environments; altered early social rearing environments through cross-fostering in Chilean flamingos is associated with limited differences in behavioural and social development.

AbstractSocial networks research using non‐human animals has grown over the past decade, utilizing a wide range of species to answer an array of pure and applied questions. Network approaches have relevance to conservation, evaluating social influences on fecundity, health, fitness and longevity. While the application of network approaches to in situ populations with conservation concern appears in published literature, the degree to which ex situ and zoo‐housed populations are the focus of “social networks for conservation research” is limited. Captive environments provide scientists with an ability to understand the social behavior of species that may be hard to observe consistently in the wild. This paper evaluates the scope of network research involving ex situ populations, analyzing output from 2010 to 2019 to determine trends in questions and subjects using ex situ populations. We show that only 8.2% of ex situ social network analysis (SNA) implications are of conservation‐focus, apparent in papers relating to birds, carnivores, bats, primates, reptiles, and ungulates. Husbandry and welfare questions predominate in ex situ network research, but over half of these papers have nonpractical application (basic science). The chance of a citation for a basic science paper was 95.4% more than for a conservation‐based paper. For taxonomic groups, primate‐focused papers had the most citations. The focus of ex situ conservation‐based networks research may be driven by the needs of conservation programs (e.g. population recovery outcomes) or by a need to evaluate the efficacy of ex situ conservation goals. We evaluate our findings considering the IUCN's One Plan Approach to conservation to show how in situ and ex situ network research is applicable to global conservation efforts. We have identified that there is a lack of application and evaluation of SNA to wildlife conservation. We highlight future areas of research in zoos and hope to stimulate discussion and collaboration between relevant parties.

Recognizing the imperative to evaluate species recovery and conservation impact, in 2012 the International Union for Conservation of Nature (IUCN) called for development of a "Green List of Species" (now the IUCN Green Status of Species). A draft Green Status framework for assessing species' progress toward recovery, published in 2018, proposed 2 separate but interlinked components: a standardized method (i.e. measurement against benchmarks of species' viability, functionality, and preimpact distribution) to determine current species recovery status (herein species recovery score) and application of that method to estimate past and potential future impacts of conservation based on 4 metrics (conservation legacy, conservation dependence, conservation gain, and recovery potential). We tested the framework with 181 species representing diverse taxa, life histories, biomes, and IUCN Red List categories (extinction risk). Based on the observed distribution of species' recovery scores, we propose the following species recovery categories: fully recovered, slightly depleted, moderately depleted, largely depleted, critically depleted, extinct in the wild, and indeterminate. Fifty-nine percent of tested species were considered largely or critically depleted. Although there was a negative relationship between extinction risk and species recovery score, variation was considerable. Some species in lower risk categories were assessed as farther from recovery than those at higher risk. This emphasizes that species recovery is conceptually different from extinction risk and reinforces the utility of the IUCN Green Status of Species to more fully understand species conservation status. Although extinction risk did not predict conservation legacy, conservation dependence, or conservation gain, it was positively correlated with recovery potential. Only 1.7% of tested species were categorized as zero across all 4 of these conservation impact metrics, indicating that conservation has, or will, play a role in improving or maintaining species status for the vast majority of these species. Based on our results, we devised an updated assessment framework that introduces the option of using a dynamic baseline to assess future impacts of conservation over the short term to avoid misleading results which were generated in a small number of cases, and redefines short term as 10 years to better align with conservation planning. These changes are reflected in the IUCN Green Status of Species Standard.

AbstractZoo‐housed pelicans are commonplace, but their breeding record is poor and little research is published on the activity patterns, as potential predictors of nesting, of captive flocks. Existing literature shows that comparative research can provide useful information for husbandry and conservation planning for pelican populations. The opportunity arose to investigate the time‐activity budget and social network of a breeding flock of captive great white pelicans. Three chicks were hatched in June and July 2016 and one in March 2017. Data on state behaviors, space use, and association preferences were collected around these nesting events, from October 2016 to February 2017 and July to October 2017. Results suggest that pre‐nesting periods were associated with heightened flock‐wide vigilance, suggesting that vigilance may be a precursor for courtship or nesting activity. Social network analysis revealed nonrandom associations between birds and a social structure across the flock, in which subadults seemed to associate more with each other than with adult birds. A limited visitor effect was noted; whilst no overall behavior change was apparent with different numbers of visitors, pelicans did widen their enclosure usage with increased visitor presence. These data are relevant to those attempting to breed this pelican, who wish to know more about the daily behavior patterns of this species across the season and physiological state, and who wish to understand pelican social structure, which is useful to the planning and implementation of bird moves or changes to the social environment of the flock. Further extending such research to include uninterrupted observation over a successful breeding event is recommended.

AbstractThe sound environment of a zoo animal is a complex milieu of animal and human‐generated sounds; coming from the species itself, other species, visitors, keepers and other zoo‐users. Research determining how different components of the sound environment affect animal behaviour is surprisingly lacking but could have real‐world impacts for animal welfare and zoo enclosure design. The current study investigated the effects of the sound environment on two flocks of flamingos housed in open‐air enclosures at British zoos. Measures of how each flock used its enclosure (as a response variable) and environmental variables (Inband Power and Peak Frequency were recorded as characteristics of the sound environment, as well as temperature, humidity and cloud cover, and finally visitor presence—all as potential predictor variables) were made over a 2‐month period. Assessment of space use by zoo animals is often used as a measure of the appropriateness of an exhibit and to understand welfare. Given that flamingo activity is influenced by weather and that the sound environment of the zoo is likely to be influenced by the number and the presence of visitors, it was assumed that these predictor variables would influence where the flamingos were located at different times of the day. As expected, there was a complicated relationship between enclosure use and Inband Power (average spectral density, a measure of sound energy) in both flocks; visitors generated salient sound but other visitor characteristics such as their physical presence may have impacted the movement of the birds around their enclosures. Results show a complex picture where environmental conditions influence flamingo enclosure usage as well as visitor presence and sounds around/in the enclosure. Findings are not consistent between the two flocks, with one flock demonstrating distinct temporal change to enclosure zone occupancy and the other responsive to humidity and cloud cover variation. We believe enclosure use can provide a valuable indication of how birds react to their soundscape; however, our findings suggest more work is needed to unpick the components of captive sound environments, and their relative effects on how animals use their space.

Flamingos are well known for their gregarious habits and aggregations in large flocks, but evaluation of the mechanisms behind social grouping remain poorly understood. Captive birds provide a useful model for investigating aspects of social choice in highly gregarious, long-lived species. Animals invest in social relationships that convey fitness benefits and bonds can be long-lasting. For some species, field-based measurement of social networks can be difficult. Captive populations therefore provide a useful alternative for measuring social choices. Data were collected on flamingos at WWT Slimbridge Wetland Centre from 2013 to 2016 and compared to data from 2012. For three flocks, associations were analysed along with individual foot health scores to identify any relationship between health and social behaviour. Long-term partnerships were present in all flocks; preferred associates noted in 2012 were present in 2016. Matrix correlations across years were positive; arrangements of dyads, trios and quartets with higher ties strengths were visible at the beginning and end of the study. Both male-male and female-female bonds were stable over time. All flamingos were more frequently seen socialising than solitary; those in the largest flock showed the highest occurrence of social behaviour (irrespective of enclosure size differences). The number of connections realised from all available within a network was significantly influenced by season. Foot health did not predict associations in these three flamingo networks. Our results indicate that flamingo societies are complex (i.e. formed of long-standing preferential partnerships and not loose, random connections) and the impact of flock size and environment on sociality should be investigated further. These results are helpful for those working with captive flamingos to consider the number of birds housed so that an array of opportunities for choice of associate and/or breeding partner are available in zoo-housed flocks.

Social network analysis (SNA) is useful for evaluating management zoo regimes to ensure that any fitness benefits of sociality are preserved in captive-housed groups. This paper explores the association patterns of 13 giraffes housed at Longleat Safari Park, UK. Wild giraffes exhibit a fission–fusion social system with preferential bonding. As zoo-housed giraffes are common, they are excellent study subjects for using SNA to investigate key aspects of sociality within a managed social environment. Social bonds were assessed over different seasons and data from two study periods (2011 and 2015) were analyzed to see consistency of “social type” (i.e. more social or more solitary). Associations showed the occurrence of consistent preferential bonds between named individuals but time of year influenced the patterns of social bonds. Female-female bonds and female-offspring bonds appeared to be strongest. For animals present in 2011 and 2015, differences in time spent socializing between years were apparent. Results suggest that giraffes may be flexible in their choice of social partner and zoo-managed herds should include a range of individuals from which each animal can choose a preferred associate.

Lechwe are specialised wetland antelope that can have a strict social hierarchy or perform lekking during breeding. The southern lechwe (Kobus leche) and the Nile lechwe (K. megaceros) are both found in zoos globally, but little research is available to support husbandry decisions. The aim of this research was to investigate current housing and husbandry used for these lechwe across North American and European zoos. A survey was distributed to holders in 2018 and information on 33 herds (18 Nile and 15 southern) was collected. The survey focussed on population demographics, enclosure size, biologically relevant exhibit features, mixed-species holdings, nutrition, use of environmental enrichment and performance of abnormal repetitive behaviours. Results showed that lechwe were housed in herds with similar sex ratios to wild counterparts but with a potential lack of opportunity to lek. Many zoos provided wetland, but this was rarely actively managed, and not all zoos provided cover for hiding and retreat. Current feeding practice differed significantly compared to available antelope husbandry guidelines. No consistency in amounts of pellet, forage or produce provided to lechwe across institutions was found. Abnormal repetitive behaviour was noted by several zoos, but no significant predictor of such behaviour could be identified. Despite some identifiable recognition of ecology informing lechwe management, it is important that evidence-based husbandry decisions are made based on a species’ evolutionary pathway and ecological needs and some fundamental features of lechwe husbandry do not always correlate with the adaptive traits of a specialised wetland ungulate.

Welfare assessment is a tool to both identify welfare challenges and to evidence where current husbandry practices support positive welfare outcomes. Such tools are becoming more available and can be amended based on the nature of the facility and needs of taxonomic groups. Currently, welfare assessment has a strong mammalian theme, and some behavioural measures of welfare commonly applied to mammals do not translate well for other taxa. This paper provides a method for welfare assessment of Anseriformes; widely housed, diverse bird species kept under a range of management styles. A mixture of resource-based (i.e. determination of aspects of the physical environment or the bird’s physical appearance or activity) and animal-based (i.e. observations that equate to a bird’s feelings or personality characteristics) measures are integrated to enable a full review of potential predictors of welfare. The method provides a rapid and valid way for all personnel to collect information that evaluates quality-of-life experiences of the Anseriformes under their care. Explanations of key terminology are provided to enable repeatable and reliable assessment for all persons using the tool. Suggestions for follow-up actions are provided to emphasise why the welfare assessment process needs to be one of continual re-evaluation of animal care.

For specialised feeders, accessing food resources may impact on the performance of appetitive foraging and social behaviours at individual and population levels. Flamingos are excellent examples of social species with complex, species‐specific feeding strategies. As attainment of coloured plumage depends upon intake of dietary carotenoids, and as study of free‐ranging flamingos shows that foraging is disrupted by aggression from other birds, we investigated the effect of four feeding styles on foraging and aggression in captive lesser flamingos. We evaluated individual and group differences in foraging and aggression when birds consumed bespoke “flamingo pellet” from a bowl, an indoor feeding pool and an outdoor feeding section of their pool. Natural foraging (when birds were feeding irrespective of the presence of pellet) was recorded for comparison with artificial feeding styles. One‐minute long video footage of the birds' activities in these different locations, recorded between 2013 and 2016, was used to evaluate behaviour. Total number of seconds engaged in feeding and in aggression was recorded by continuous sampling. The colour of individual birds was scored from 1 (mainly white) to 4 (mainly pink). For natural filter feeding in the outdoor pool, maximum foraging was twice as much as bowl feeding, whilst aggression was less than half as much as other feeding methods. Overall, a more restricted feeding style significantly predicted aggression, along with increasing group size. Plumage colour significantly influenced aggression (brightest flamingos were more aggressive) and showed a non‐significant trend with foraging (brighter birds fed less than paler birds). No sex effect on feeding or aggression was found. This study enhances our understanding of husbandry and species' biology impacts on captive behaviour and provides data‐based evidence to improve food presentation. For flamingos, implementation of spacious outdoor feeding areas can encourage natural foraging patterns by reducing excess aggression and enhances welfare by improving flock social stability.

Same-sex pair bonds have been documented in several animal species and they are widespread in birds. However, little is known about the evolutionary origin and the adaptive value of such behaviour. The aim of this study was to investigate the parental behaviour of four zoo female greater flamingos involved in two breeding pairs, housed in a flock at Parco Natura Viva, Italy. Further, the behaviour of the study females was compared with that of male and female flamingos in heterosexual pairs described in a previous published work on this same flock. For each pair, the behaviour of both birds during the incubation period was recorded and twenty 10-minute sessions were run within the incubation period. A continuous focal animal sampling method was used to collect data on location (on the nest or not on the nest) and the parental care behaviour (e.g.: agonistic behaviours toward disturbing conspecifics, egg-care, nest-building, self-comfort behaviour, sleeping) of the two pairs. Data of the current study females were compared with those of females and males involved in heterosexual pairs of this same flock. Results showed that within each pair the egg-layer female stayed away from the nest more than the other female. In addition, the female that did not lay an egg was more involved in agonistic behaviour compared to other females, particularly when in specific locations. In heterosexual pairs, male flamingos were more involved in the incubation and in nest protection. Moreover, no significant differences in the time spent on the nest and away from the nest between the heterosexual male and the non-layer females of same-sex pairs were found. The same findings were reported when comparing heterosexual females and the egg-layer females of the same-sex pairs. Therefore, our findings suggest that in greater flamingos the behaviour of the female–female pairs seems to be equivalent to that of male-female bonds. Such research provides more insight into flamingo social behaviour, and their reproductive cycle, and provides information on why pair bonds may form and how these affect the wider breeding behaviour of the flock.

Turacos (Musophagidae) are common zoo birds; the 14 species of Tauraco being most often exhibited. Turacos possess unique non-structural, copper-based feather pigments, and a specialized dietary strategy. Tauraco inhabit tropical woodlands, foraging for predominantly folivorous and/or frugivorous food items. Using a study population of 16 red-crested turacos (T. erythrolophus) at seven zoos in the United Kingdom, the nutrient composition of diets from diet sheets was calculated, using Zootrition v.2.6, Saint Louis Zoo, USA for analyses of important nutrients within each diet, and compared against an example of currently available literature. For all nutrients analyzed, significant differences were noted between amounts presented in each zoo's diet (as fed). Turacos are presented with a wide range of ingredients in diets fed, and all zoos use domestic fruits to a large extent in captive diets. Similarities exist between zoos when comparing amounts of as-fed fiber. Analysis of the calcium to phosphorous ratio for these diets showed there to be no significant difference from the published ratio available. While this is a small-scale study on only a limited number of zoos, it provides useful information on current feeding practice for a commonly-housed species of bird and highlights potential areas of deviation away from standard practice, as well as identifying ways of reducing wastage of food. Data on wild foraging behavior and food selection, or collaboration with tauraco keepers from institutions in the tropics, is recommended as a way of improving feeding regimes and updating feeding practice for this and other Tauraco species.

In this response, we show how the points raised in Rose et al. (2017) are relevant across taxa. The original article reviewed literature on 3 basic “groups” of animal, with a specific remit of identifying welfare needs within these groups. The focus of this article did not intend to exclude other types of animal but to show the extent of research needs in those already studied. The ideas presented are relevant to those studying other taxa, specifically scientists and zoo biologists with more expertise and knowledge of invertebrates. We feel that there is much to be gained from collaboration between individuals and institutions to adapt the questions for assaying welfare proposed by Rose et al. (2017) to a wider range of captive vertebrate and invertebrate taxa.

Aggressive behavioural interactions between animals are widespread in nature, with ecological and evolutionary consequences of such interactions reported for both individuals giving and receiving aggression. Yet despite the importance of aggressive interactions in determining social dominance and conferring fitness benefits to successful individuals, we lack a general understanding of the conditions that influence the variation in agonistic behaviours among individuals and species. We conducted a global meta-analysis of published time–activity budgets, using a data set comprising 555 values from 88 studies, to determine variation in the time that waterbirds engaged in aggressive interactions. The mean ± SD percentage of time devoted to aggression was 2.0 ± 3.6%, with a range of 0.0–35.0%. We used our data set to test four predictions regarding avian aggression, based on the findings of earlier site-based studies. We predicted that the time spent on aggression would be influenced by four factors: (1) age class, (2) sex, (3) seasonal timing and (4) functional feeding group. A comparison of linear mixed-effects models using an information theoretic approach indicated that the proportion of time waterbirds spent engaged in aggressive interactions was best explained by the age class and sex of the focal waterbird species. More time was spent on aggression by males than females, and by adults than juveniles. We found no evidence that the time spent on aggressive behaviours varied across latitudes or body mass, with seasonal timing, sexual dimorphism, migration or breeding strategies, or between different functional feeding groups. Our findings highlight the high levels of variation in the time devoted to aggression across species, feeding groups, latitudes and seasons. Furthermore, our study demonstrates the utility of the numerous published time–activity budget studies that are available as a valuable source of data that can be used to answer broad-scale questions regarding animal behaviour.

Flamingos are ubiquitous captive species (potentially the world's most commonly kept zoo bird) that have long lifespans and unique breeding cycles. In-depth research into the links between provision (enclosure, husbandry), behavioural performance (reflecting internal motivation) and perceived welfare state (from behavioural cues) can inform management for good welfare over the many decades of a bird's life, and benefit reproductive output. Here, the published literature on flamingo husbandry is reviewed, with reference to our current understanding of flamingo behaviour in the wild. Evaluation of whole-flock time budgets and assessment of behavioural diversity can highlight any deviation from a norm. Several published works suggest ways of improving breeding success in captive flamingos by re-evaluating husbandry routines as well as highlighting minimum numbers of birds per flock for 'good welfare'. Research has shown that some aspects of zoo-flamingo activity can match that of wild birds. Further examination of specific behaviour patterns, as well as the motivations for these, would allow for evidence-based enclosure design and provision of species-specific behavioural husbandry. Future research topics covering social support, foraging activity and developing a definition of 'positive welfare activity' would further enhance zoo management practices for these birds. © 2014 the Zoological Society of London.

The Malayan, or Asian, tapir (Tapirus indicus) has a diminishing wild population and is becoming more common in captivity as zoos attempt to manage sustainable ex situ populations. Tapirs can be relatively easy to maintain and breed, but captive animals appear to suffer from reduced activity budgets, obesity, and poor public image. A questionnaire-based survey was designed and sent specifically to 10 collections around the world that exhibit Malayan tapirs, with the aim of assessing husbandry regimes to determine prevalence of standardized practices as well as highlighting any key differences, and to showcase good practice, thus providing information beneficial to those maintaining this species in their zoo. Twenty-five animals were included in the survey from collections across four continents. The research's major conclusions show differing dietary make-up, with a lack of forage provision, contrasting with a diverse array of enrichment protocols used. Significant differences were noted between zoos for total amount of food offered (P = 0.000) as well as ratios of forage to concentrate pellet offered (P = 0.004). Comparing food offered to male and female tapirs with published requirements for an "average" of either gender shows not all zoos providing the amount suggested in husbandry guidelines. Intelligently designed and original enrichment was provided to all animals but differences between zoos were noted in the application and "usefulness" of enrichment for individual tapir. Overall, animals are benefiting from enrichment but welfare could be further improved via consistent feeding of ad libitum forage and regular use of browse as a constituent part of daily rations. © 2012 Wiley Periodicals, Inc.