Engaging internal and external audiences to develop and promote zoo-based conservation efforts.

As place-based conservation organizations, zoos are in a central position to support individuals in making small changes in their lives that will support the protection of wildlife and their habitats. This paper describes the secondary analysis of data collected from multi-phase front-end, exploratory evaluation that informed the development of a conservation action campaign in association with a non-profit, urban zoo. In phase one, internal organization staff were invited to attend workshops during which they brainstormed potential conservation actions that they felt were important for the zoo to promote. They identified and ranked 164 unique actions. In phase two, the ranking was used to narrow down the 164 actions to 20 actions which were used to develop a survey administered to visitors who opted in to receiving online surveys from the zoo. The survey asked participants to state their interest in each of the 20 conservation actions. The Transtheoretical Model of Behavior Change informed the analysis of responses. Through this approach we identified actions that people were already doing, interested in doing, and not interested in doing. The responses from this survey were used to narrow down the list further to 10 actions used in a survey in phase three. This second survey administered to zoo visitors on grounds asked participants which of the 10 actions they would be most interested in doing, and the perceived barriers and benefits of doing them. This process allowed us to use evidence-based decision making to choose which conservation actions would resonate most with the community for our conservation action campaign. We also were able to identify values visitors held that might influence environmentally friendly behaviors. Visitors who responded to this survey tended to respond in ways that aligned with self-transcendent values. The research suggests that the campaign should focus on habitat restoration and remediation and purchasing wildlife friendly coffee and other products.

SELECTED INSTANCES OF ELEPHANT ENDOTHELIOTROPIC HERPESVIRUS SHEDDING IN TRUNK SECRETIONS BY AFRICAN ELEPHANTS (LOXODONTA AFRICANA) IN COMPARISON TO SHEDDING BY ASIAN ELEPHANTS (ELEPHAS MAXIMUS).

This study examined the viral shedding kinetics of elephant endotheliotropic herpesvirus (EEHV) in African elephants (Loxodonta africana) compared to viral shedding behavior in Asian elephants (Elephas maximus). Little is known about the transmission dynamics and epidemiology of this disease in African elephants. In light of recent clinical cases and mortalities, this paper aims to identify trends in viral biology. Trunk wash samples were collected from 22 African elephants from four North American zoological institutions that had recently experienced herd viremias or translocations. Processing of these samples included DNA extraction followed by qPCR to quantitate viral DNA load. The results were then compared with available literature that chronicled similar cases in Asian and African elephants. Minimal EEHV shedding was detected in response to varied herd translocations. Increased shedding was recorded in herds in which an elephant experienced an EEHV viremia when compared to baseline shedding. These index infections were followed by subsequent viremias in other elephants, although it is not known if these were recrudescence, transient controlled viremias, and/or primary infections via transmission to other elephants. When compared to historically published data, it was observed that EEHV3 cases in African elephants and EEHV1A cases in Asian elephants had consistently higher levels of viral DNA in the blood than were shed in trunk secretions, a fact that is seemingly inconsistent with such severe cases of disease and the high mortality rates associated with those respective types. The findings produced in this study highlight the need for more routine monitoring of viral shedding in African elephant herds to elucidate possible EEHV transmission and recrudescence factors for ex situ population management.

Positive clinical experience with paromomycin sulfate in treatment of Balantioides coli (=Balantidium coli) natural infection in zoo-kept Mandrill monkeys (Mandrillus sphinx) and Western lowland gorillas (Gorilla gorilla gorilla).

Balantioides coli (=Balantidium coli), a large ciliated protozoan, is reported in multiple free-ranging and captive primate species, often in association with a clinical presentation that requires medical intervention. This report describes the clinical effectiveness of paromomycin sulfate against B.coli in zoo-kept mandrill monkeys (Mandrillus sphinx, at orally doses of 8-31 mg/kg, once daily (SID) for 7 days) and gorillas (Gorilla gorilla gorilla, at orally doses of 1.4-3.1 mg/kg, SID for 5 days).

A SYSTEMATIC REVIEW OF RETROSPECTIVE MORBIDITY AND MORTALITY STUDIES ON CAPTIVE WILDLIFE SPECIES.

Zoological institutions manage animals for conservation, education, entertainment, and research purposes. Zoological staff have a responsibility to safeguard the welfare of animals in their care. Retrospective morbidity and/or mortality studies (MMSs) can be useful tools to highlight common diseases in captive wildlife populations. There is currently no standardized methodology for conducting MMSs. Variation in the methodology of MMSs, particularly the categorization of diseases, can make comparisons between studies challenging and may limit the applicability of the results. A Preferred Reporting Items for Systematic Reviews and Meta-analyses (PRISMA) compliant systematic review was performed, which identified 67 MMSs describing 146 species of captive wildlife. These MMSs are becoming more common and were predominantly performed on mammals (76/146). Prospective authors are encouraged to perform MMSs on amphibians, birds, reptiles, fish, and invertebrates. The studied animals were mostly managed at institutions in the United States of America (28/67, 41.2%) and Europe (14/67, 20.9%). Classifying individuals into age groups facilitates the identification of disease trends within age classes. Only 22/67 (32.8%) studies cited justification for their age classification; classifications should be based on a referenced source on the breeding biology of the studied species. There is variation in the body systems used by authors and into which system a disease is categorized, which makes study comparisons challenging. Diseases were predominantly categorized by etiology and body system (28/77, 36.4%). Because of its ubiquity, the use of the categorization system employed by the pathology module of the Zoological Information Management System is recommended as a useful standard. This system is imperfect, and amendments to it are suggested. The results and recommendations of this study were discussed with a panel of zoo and wildlife experts; guidelines have been formulated for prospective authors aiming to conduct MMSs in captive wildlife.

Breeding behavior analysis in a large captive colony of African penguins (Spheniscus demersus) and its implications for population management and conservation.

The African penguin Spheniscus demersus, frequently housed in zoos, holds potential for future reintroduction efforts due to its declining wild population. This paper aims to explore various aspects of reproductive performance in African penguins within a large ex situ colony at Zoo Wroclaw in Poland, covering 9 years of breeding behaviors. The analysis reveals parallels in colony growth and partner change patterns with those observed in the wild. Positive correlations were found between breeding success and pair-bond duration, with the increasing colony size influencing reproductive performance. Contrary to their wild counterparts, captive African penguins initiate breeding attempt and produce a fertilized egg at a younger age. However, successful breeding still requires gaining experience or forming pairs with more experienced partners. Our research indicates that providing captive African penguins with unlimited food resources and sufficient nesting space results in rapid colony growth. The increased colony size facilitates breeding behaviors that positively influence population dynamics, particularly through the maintenance of long-term pair bond relationships and the potential for partner changes when necessary or desirable to enhance breeding success. We present compelling case studies in pair fidelity, offering valuable insights and implications for the management of captive populations and conservation efforts.

Case study: Visual barriers reduce pacing in captive tigers.

Captive large felines are prone to abnormal repetitive behaviors like pacing, which are associated with welfare issues. Visual contact without the opportunity to engage in appropriate behavior is known to increase pacing. To better understand the relationship between pacing and conspecific visual contact, we investigated this effect by conducting a barrier experiment on a male-female pair of Sumatran tigers (Panthera tigris sumatrae) in Rotterdam Zoo, the Netherlands. The tigers were exposed to four consecutive housing treatments: (i) housed in the same enclosure (baseline), (ii) housed in separate enclosures with visual contact, (iii) housed in separate enclosures without visual contact, and (iv) housed in the same enclosure after the separation. We used focal and scan sampling to measure pacing and recorded the number of visitors. Moreover, we applied scan sampling to measure activity. Overall, our results indicate that the tigers paced significantly more when housed in separate enclosures with conspecific visual contact. Moreover, our results suggest that limiting visual contact between neighboring tigers can mitigate pacing. Implementing these findings in tiger husbandry and enclosure design has the potential to improve animal welfare zoo populations of large felines.

Behavior of polar bear (Ursus maritimus) cubs post-den emergence at the Detroit Zoo.

In the wild, female polar bears (Ursus maritimus) with cubs may spend extended periods of time within the den following initial emergence. As a result, studying behavioral development of cubs at the denning site has been difficult and unreliable. Although care staff at zoological institutions have easier access to animals, every effort is made to minimize intrusions to maternal groups, which presents research limitations. Detroit Zoological Society staff used audio-equipped cameras installed in behind-the-scenes spaces to conduct approximately 9 weeks of monitoring on two female polar bear cubs of the same litter, one of whom (Laerke) was hand-reared while the other (Astra) was mother-reared. Monitoring spanned ages 14-24 weeks and consisted of 12, 5-min focal observations per observation day timed to evenly cover the entire 24-h day. Using generalized linear mixed models, we examined relationships between behavior and time of day, hour, and age. We also conducted descriptive analyses and used these to draw comparisons between the two cubs where appropriate. Despite different rearing environments, both cubs had highly comparable patterns of inactive, locomotory, and independent play behaviors. Astra generally decreased time spent nursing and in proximity to Suka as she aged. Data presented here represent a continuation of previous neonatal observations conducted on Astra, adding to the minimal body of knowledge currently available on first year polar bear cub development. We also report successful mitigation of stereotypic behavior in Laerke. Further studies of captive maternal groups can provide insight into this critical developmental stage, supporting both captive-based breeding efforts and in situ conservation efforts.

Traumatic telson avulsion and attempted external stabilization in Atlantic horseshoe crabs (Limulus polyphemus).

Atlantic horseshoe crabs (Limulus polyphemus) are prevalent in public aquarium touch pools. Despite their popularity, the literature concerning medical management under managed care is sparse. Noninfectious conditions include trauma to the exoskeleton and compound eyes; however, injury to the soft tissue connecting the telson and opisthosoma has not been previously reported. This report describes telson avulsion in three Atlantic horseshoe crabs at a public aquarium and attempted external stabilization with a vascular silicone tie in two of three affected animals. The horseshoe crab that received no veterinary intervention suffered a complete telson amputation 1 month after the initial injury. Although the two other horseshoe crabs did well postoperatively, and external stabilization prevented further avulsion or amputation, they still could not right themselves if flipped into dorsal recumbency. All three horseshoe crabs were ultimately euthanized due to animal welfare concerns. To avoid potentially serious telson ligament injury, horseshoe crabs should never be picked up by the telson. The authors recommend that telson ligament injuries be addressed promptly, any inciting cause be mitigated, and external stabilization be considered earlier.

Female long-term sperm storage results in viable offspring in the Himalayan Mountain Pitviper, Ovophis monticola.

The ability of females to store sperm for extended periods in their reproductive tracts (termed long-term sperm storage, LTSS) has been reported across a diversity of vertebrate taxa. The evolutionary, ecological, and physiological significance of LTSS is wide-ranging and includes the ability to produce offspring when mates may be temporally scarce by way of decoupling copulation from ovulation, inbreeding avoidance, and the generation and maintenance of genetic diversity in progeny. Among vertebrate lineages, nonavian reptiles exhibit a remarkable capacity for LTSS, with the production of viable offspring reported after periods exceeding 6 years since prior contact with a potential mate. Given that female reptiles are able to store viable sperm for prolonged periods, it is important to disentangle LTSS from that of facultative parthenogenesis (FP), a reproductive trait which appears widespread among all reptile lineages. The implications of this distinction are particularly important in the context of the development and management of captive breeding programs. To accurately determine between the two reproductive strategies, genomic screening is highly recommended. Following a period of isolation for 13 months from a potential male mate, a female Himalayan Mountain Pitviper (Ovophis monticola) produced a clutch of three male offspring. Here, through genome-scale analyses of the female and her progeny, we document the first record of LTSS in this genus and exclude FP as the alternative hypothesis.

First successful artificial insemination of the endangered Louisiana pinesnake, Pituophis ruthveni.

The Louisiana pinesnake (Pituophis ruthveni) is considered one of the rarest snakes in North America and was federally listed under the Endangered Species Act in 2018. Captive breeding and reintroduction of zoo-bred hatchlings has been successful, however, limited founders in the captive population and the inability to bring new, wild genes into the captive colony presents a major concern for the conservation of this species. The use of artificial insemination (AI) was first applied to snakes in the 1980s but further development of the technique has since received little attention. Our goal was to develop a method of AI for use in breeding Louisiana pinesnakes to facilitate gene flow from wild to captive populations. We inseminated two captive Louisiana pinesnakes with semen collected from one donor male, novel to both females. Timing of AI occurred following the emergence of females from brumation, and when large, distinct follicles were detected using digital palpation. Females were inseminated four and five times over a period of 14 and 19 days, respectively, using fresh and 2-day refrigerator stored semen. One female laid seven eggs, which resulted in four fertile eggs and two viable hatchlings, while the second female produced three fertile of seven eggs laid but no viable hatchlings. Genetic analyses confirmed the donor male was the sire of hatchlings. This is the first successful AI of an endangered snake species and provides a framework for the use and optimization of assisted reproductive technologies for use in conservation breeding programs.

Anesthesia and humane euthanasia methods for king worm larvae (Zophobas morio).

A growing spotlight on insect welfare as research subjects, fodder for vertebrate animals, and patients for veterinarians highlights the need to establish humane euthanasia methods for these species. An insect of increasing commercial importance is the king worm (Zophobas morio), a type of darkling beetle larvae. The initial objective of this study was to determine if prolonged exposure to isoflurane would lengthen anesthetic recovery and lead to euthanasia. Larvae were exposed to isoflurane for 10 min, 40 min, 70 min, 190 min, 13 h, and 24 h (n = 10 for each treatment duration group), and time to recovery was subsequently monitored in room air. The second objective was to determine effectiveness of secondary euthanasia methods: injectable potassium chloride at 1 mg/g body weight (n = 5) and 3 mg/g body weight (n = 5) doses; immersion in 200 proof ethanol (n = 10), 10% buffered formalin (n = 10), and water (n = 10); and freezing at -18°C (n = 10). Prolonged exposure to isoflurane led to extended recovery times, but 100% mortality was not achieved. Submersion in ethanol and injection of potassium chloride at a 3 mg/g body weight dose resulted in 100% mortality. If these larvae are to be used as future food sources, a humane euthanasia method that preserves the ability to be consumed is required.

The effects of preformed vitamin A and provitamin A carotenoid supplementation on tadpoles of the poison frog Phyllobates vittatus.

Understanding the nutritional requirements of captive animals is necessary for proper animal husbandry, however, the specific dietary requirements for many amphibian species commonly kept in captivity are unknown. Like most vertebrates, frogs cannot synthesize carotenoids and must therefore obtain these essential nutrients through diet. It is unclear if amphibians can cleave provitamin A carotenoids to form vitamin A metabolically within the body, so common practice is to supplement their captive diets with both preformed vitamin A and provitamin A carotenoids. We carried out a feeding experiment in tadpoles of Phyllobates vittatus, a commonly kept poison frog species, to test the effects of supplementing a fish flake diet with a provitamin A carotenoid (2.5 mg/g ß-carotene) and vitamin A (0.033-0.066 µg/mL retinyl acetate), both individually and in combination. Contrary to our expectations, supplementation had either no effect or adverse effects on tadpole growth and survivorship. Tadpoles reared under supplemented diets with vitamin A showed higher mortality rates, coupled with symptoms of hypervitaminosis A. Survivors had a smaller body size and mass at metamorphosis. ß-carotene supplementation alone had no detectable effect. The vitamin A and ß-carotene levels in our supplemented diet have been shown to be harmless or benefit tadpoles of other species, yet our results indicate that adding these amounts to what is found in a generalist fish flake mix can have detrimental effects on P. vittatus tadpoles. More broadly, this study highlights the importance of creating husbandry guidelines based on the specific physiological needs of the species (or species groups) being kept in captivity, rather than general ones for all amphibians, as is often done.

A perspective on ungulate management and welfare assessment across the traditional zoo to large landscape spectrum.

Wild ungulates are managed in human care in a range of settings from traditional zoos to large ranches. These varied settings present different portfolios of risks for good or poor welfare, which leads some to question whether a particular setting is "good for welfare" and have frustrated others interested in comparing the welfare of ungulates across these settings. Differing housing and management scenarios present different challenges and opportunities in terms of welfare but this commentary posits that good welfare is possible in all of these settings. In this commentary, we also consider natural behaviors that may, at face value, compromise welfare and discuss how taking a long view on welfare addresses concerns about these behaviors, in part using arguments related to normal behavioral development that likely improves welfare at other life stages. We also highlight the role of motivation in seemingly welfare-compromising behaviors. Finally, some indicators of welfare that we believe transcend management scenarios, and are thus able to be compared across scenarios, are suggested.

Using qualitative behavior assessment to investigate the effect of tourist presence on the welfare in captive tigers (Panthera tigris) in three tourism facilities in Thailand.

Numerous facilities around the world offer tourists interactive experiences with captive tigers. Yet, the animal welfare implications of this practice have not been widely studied. This study aimed to investigate whether qualitative behavioral assessment (QBA) could: (i) provide a valid indicator of tiger's emotional state and (ii) be applied to assess whether unfamiliar human presence with hand-raised captive tigers had an impact on the emotional state of those tigers. To investigate this, QBA was applied to video clips of hand-raised captive tigers from three sites (two offering unfamiliar human interaction, Sites A and C, and one retirement site with no direct interactions, Site B) in Thailand. QBA allows inferences to be made about animal emotion on the basis of descriptions of behavioral expression. Analysis, using a free choice profiling methodology, was provided by observers (N = 38) split between three groups; tiger keepers and vets from the Thai venues (n = 12), UK-based animal behavior MSc and vet students (n = 16), and international tiger keepers (n = 10). Tigers (N = 35) were split between Sites A (n = 7), B (n = 18), and C (n = 10) and filmed at three time points; morning (0800-0930 h); midday, (1130-1230 h); and evening, (1630-1830h) totaling 105 clips. Using generalized procrustes analysis, a consensus profile was calculated for each observer group. Two meaningful dimensions of behavioral expression, explaining 75.0% of the variation, were observed across these groups: Dimension 1 (D1: "active"/"interested"/"agitated" to "relaxed"/"calm"/"chilled-out") and Dimension 2 (D2: "bored"/"stressed"/"frustrated" to "relaxed"/"curious"/"interested"). There was clear agreement between the three observer groups in terms of tiger emotional expression along D1. However, agreement was more variable on D2. The behavioral expression on D1 was not significantly affected by site but was significantly affected by an interaction between age and time of day. Time of day also affected scores on D2, with the Thai observer group also showing an effect of site. During the midday period, when unfamiliar humans were present, all tiger age groups showed more positive behavioral expressions on D1 (lower scores: "relaxed"/"calm"/"chilled-out") and more negative behavioral expressions on D2 (higher scores: "bored"/"stressed"/"frustrated"), which could indicate that the presence of unfamiliar humans was a stressor. However, tigers in the retirement Site C also displayed similar behavioral expressions, which could indicate a deeper welfare issue. With further development, QBA could be used as part of a valid tool for long-term measurement of behavioral expression in captive tigers.

A cross-over dietary intervention in captive cheetahs (Acinonyx jubatus): Investigating the effects of glycine supplementation on blood parameters.

Captive cheetahs are prone to unusual diseases which may be attributed to their high muscle meat, collagen deficient captive diet. Glycine is a simple amino acid that is abundant in collagen rich tissues and has many physiological functions, specifically in collagen synthesis and in the conjugation of detrimental by-products produced during gut bacterial fermentation. Therefore, the aim of this study was to investigate the effect of a 4 week glycine supplementation on the body measurements, haematology and serum blood parameters of 10 captive cheetahs using a randomised controlled cross-over design. This approach has not yet been used to investigate the effect of diet in captive cheetahs. Cheetahs were randomly assigned to a control diet (horse meat only) or a glycine diet (30 g glycine per 1 kg meat) for 4 weeks before being crossed over. Blood was collected at baseline and after each intervention. The glycine diet resulted in a decreased serum albumin, alkaline phosphatase and total calcium concentration and increases in eosinophils and basophils counts compared to the control diet. Body weight also decreased on the glycine diet which may be due to increased ß-oxidation and fat loss. This was the first study to investigate the effect of glycine supplementation, which resulted in slight body and blood changes, in captive cheetahs using a cross-over design and this approach should be utilised for future dietary studies.

UVB-emitting LEDs for reptile lighting: Identifying the risks of nonsolar UV spectra.

UVB lamps are used to provide reptiles housed indoors with the UV radiation necessary to synthesize vitamin D3 in their skin. Since 2019, UVB-LED lamps have been on sale for use in reptile husbandry. We performed spectral analysis and mapped the UV irradiance for 18 of these lamps. The positive benefits of UVB-LED lamps over traditional products include greater energy efficiency, freedom from mercury and easy installation without external ballasts. However, the spectra of all the UVB-LED lamps tested had little similarity to the solar UV spectrum. Some lamps emitted short-wavelength, non-terrestrial, radiation known to cause acute photo-kerato-conjunctivitis; we report one case. All lamps were lacking significant output in the range 315-335 nm, essential for natural self-regulation of cutaneous vitamin D3 synthesis, preventing overproduction. We describe a possible risk of serious hypervitaminosis D based on our spectral analysis. We call for long-term animal studies to assess this risk, in which the reptiles under these lamps are exposed to species-appropriate UV index levels according to their Ferguson Zone allocation and serum levels of vitamin D3 and 25(OH)D3 monitored. Spectral modifications of the lamps to make the spectrum more like sunlight may be an essential way of mitigating this risk.

Never be mute about bird welfare: Swanning around with environmental enrichment.

Environmental 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.

An evaluation of DNA sample source and molecular markers to determine gender in the short-beaked echidna (Tachyglossus aculeatus).

The short-beaked echidna is sexually monomorphic such that gender identification without veterinary intervention is challenging. The aim of this study was to evaluate and compare the most optimal noninvasive genetic source by extracting echidna genomic DNA (gDNA) from fecal scats, plucked hair, and quills to perform genetic sex testing using a range of molecular markers. Sex determination of 14 captive short-beaked echidnas was determined by amplifying isolated DNA from noninvasive samples, targeting two Y chromosome (male-specific) genes (mediator complex subunit 26 Y-gametologue [CRSPY] and anti-Müllerian hormone Y-gametologue [AMHY]), in addition to four confirmed sex-specific RADseq markers. Results of noninvasive samples were compared with blood samples and clinical records. Receiver operating characteristic curves were used to assess accuracy of sex determination of markers for each sample type. The gender of the echidnas was successfully identified on 75% of occasions using fecal samples, 90.6% occasions using hair, and 84.6% occasions with quills. Overall, the male-specific RADseq markers accurately identified the sex of echidnas with all sample types for 90% of animals; compared with 81.5% using CRSPY, and 82.0% using AMHY to identify sex. Collection of hair, quills, and feces provides a useful alternative to invasively collected samples, however, the accuracy of results depends on sample type and genetic marker selected. We found gender determination in the short-beaked echidna was most accurate using four male-specific RADseq markers on gDNA isolated from blood and hair. The noninvasive genetic sexing techniques documented here will inform and facilitate husbandry and genetic management of captive echidna populations.

Effect of four eggshell repair materials on weight loss during incubation of white leghorn chicken eggs.

Egg weight loss during incubation is a key indicator used to monitor successful egg development and is closely related to hatchability and chick survival. Artificial incubation is one of the most important captive breeding techniques used in conservation efforts to bolster avian populations. To repair damage to the eggshell and ensure embryonic viability during incubation, a variety of repair coverings can be applied. This study tested the impact of four repair materials (nail polish, synthetic glue, medical dressing, and molten wax film) on egg weight loss during incubation. We found no impact on weight loss for coverings smaller than 35% of the eggshell surface, nor did we find any differences between covering types. The average egg weight loss decreased as the coverage area increased, and the weight loss did not differ when blunt versus sharp-end coverings were compared. Given the relative insensitivity of egg weight loss and survival to the type of patch material used, we concluded that the selection of material for the purpose of weight loss management could be based on practical considerations, such as ease of application and availability.

Putting zoo animal cancer into perspective.

As part of a comparative research agenda that promises insights that help extend the human lifespan and combat cancer, cancer prevalence in zoo animals has received recent attention. Here, we want to draw attention to a principle of cancer research that was introduced into the zoo world as early on as 1933, but that seems to have gone somewhat forgotten: Cancer is mainly a disease of old age, and therefore studies aiming at identifying taxa that are particularly susceptible or resistant to cancer must control for whether the respective zoo populations are 'old.' In a comparative context, 'old age' cannot be measured in absolute terms (e.g., years), but only in relation to a species' maximum lifespan: Species that achieve, across zoos, a higher mean lifespan as a percent of their maximum lifespan are 'older.' When applying this metric to former as well as more recently published data on cancer prevalence, it appears that those species that become relatively old in zoos-in particular, the carnivores-have a relatively high cancer prevalence. Any improvement in animal husbandry-which reduces premature deaths-should, by default, lead to more cancer. Cancer in zoo animals, like any other old-age condition, might therefore be embraced as a proxy for good husbandry. Rather than following a sensationalist approach that dramatizes disease and death per se, zoos should be clear about what their husbandry goals are, what relative longevities they want to achieve for which species, and what old-age diseases they should therefore expect: in the end, one has to die of something.