Comparative mortality levels among selected species of captive animals

We present life tables by single year of age and sex for groups of animals and for 42 individual mostly mammalian species. Data are derived from the International Species Information System. The survivorship of most of these species has never been mapped systematically. We demonstrate that, in most of the groups, female survivorship significantly exceeds that of males above age five. Wild-born animals do not have mortality that differs significantly from captive-born animals. While most species have mortality that rises with age above the juvenile stage, there are several groups for which the age pattern of mortality is nearly level.ISIS, longevity, mortality, survivorship

Comparing life expectancy of three deer species between captive and wild populations

Life in zoological gardens provides a number of benefits to captive animals, resulting in an artificial reduction of the "struggle for life” compared to their free-ranging counterparts. These advantages should result in a higher chance of surviving from 1year to the next, and thus in longer average life expectancies for captive animals, given that the biological requirements of the species are adequately met. Here, we compare the life expectancy of captive and free-ranging populations of three deer species (reindeer Rangifer tarandus, red deer Cervus elaphus, and roe deer Capreolus capreolus). Whereas captive reindeer and red deer had life expectancies equal to or longer than free-ranging individuals, the life expectancy of captive roe deer was shorter than that of free-ranging animals. These results support the impression that roe deer are difficult to keep in zoos, whereas reindeer and red deer perform well under human care. We suggest that the mean life expectancy of captive populations relative to that of corresponding free-ranging populations is a reliable indicator to evaluate the husbandry success of a species in captivit

Basic considerations on seasonal breeding in mammals including their testing by comparing natural habitats and zoos

Seasonal reproduction is common in mammals. Whereas specific conditions triggering a seasonal response can only be identified in controlled experiments, large-scale comparisons of reproduction in natural habitats and zoos can advance knowledge for taxa unavailable for experimentation. We outline how such a comparison can identify species whose seasonal physiology is linked to photoperiodic triggers, and those whose perceived seasonality in the wild is the consequence of fluctuating resources without a photoperiodic trigger. This concept groups species into those that do not change their aseasonal pattern between natural habitats and zoos because they are not constrained by resources in the wild, those that do not change a seasonal pattern between natural habitats and zoos because they are triggered by photoperiod irrespective of resources, and those that change from a more seasonal pattern in the natural habitat to an aseasonal pattern in zoos because the zoo environment alleviates resource limitations experienced in the wild. We explain how detailed comparisons of mating season timing in both environments can provide clues whether a specific daylength or a specific number of days after an equinox or solstice is the likely phototrigger for a taxon. We outline relationships between life history strategies and seasonality, with special focus on relative shortening of gestation periods in more seasonal mammals. Irrespective of whether such shortening results from the adaptive value of fitting a reproductive cycle within one seasonal cycle (minimizing ‘lost opportunity’), or from benefits deriving from separating birth and mating (to optimize resource use, or to reduce infanticide), reproductive seasonality may emerge as a relevant driver of life history acceleration. Comparisons of data from natural habitats and zoos will facilitate testing some of the resulting hypotheses

Comparing life expectancy of three deer species between captive and wild populations

Life in zoological gardens provides a number of benefits to captive animals, resulting in an artificial reduction of the “struggle for life” compared to their free-ranging counterparts. These advantages should result in a higher chance of surviving from one year to the next, and thus in longer average life expectancies for captive animals, given that the biological requirements of the species are adequately met. Here, we compare the life expectancy of captive and free-ranging populations of three deer species (reindeer Rangifer tarandus, red deer Cervus elaphus, and roe deer Capreolus capreolus). Whereas captive reindeer and red deer had life expectancies equal to or longer than free-ranging individuals, the life expectancy of captive roe deer was shorter than that of free-ranging animals. These results support the impression that roe deer are difficult to keep in zoos, whereas reindeer and red deer perform well under human care. We suggest that the mean life expectancy of captive populations relative to that of corresponding free-ranging populations is a reliable indicator to evaluate the husbandry success of a species in captivity

Historical development of the survivorship of zoo rhinoceroses—A comparative historical analysis

Zoo animal husbandry is a skill that should be developing constantly. In theory, this should lead to an improvement of zoo animal survivorship over time. Additionally, it has been suggested that species that are at a comparatively higher risk of extinction in their natural habitats (in situ) might also be more difficult to keep under zoo conditions (ex situ). Here, we assessed these questions for three zoo‐managed rhinoceros species with different extinction risk status allocated by the IUCN: the “critically endangered” black rhino (Diceros bicornis), the “vulnerable” greater one‐horned (GOH) rhino (Rhinoceros unicornis), and the “near threatened” white rhino (Ceratotherium simum). Comparing zoo animals ≥1 year of age, the black rhino had the lowest and the white rhino the highest survivorship, in congruence with their extinction risk status. Historically, the survivorship of both black and white rhino in zoos improved significantly over time, whereas that of GOH rhino stagnated. Juvenile mortality was generally low and decreased even further in black and white rhinos over time. Together with the development of population pyramids, this shows increasing competence of the global zoo community to sustain all three species. Compared to the continuously expanding zoo population of GOH and white rhinos, the zoo‐managed black rhino population has stagnated in numbers in recent years. Zoos do not only contribute to conservation by propagating ex situ populations, but also by increasing species‐specific husbandry skills. We recommend detailed research to understand specific factors responsible for the stagnation but also the general improvement of survivorship of zoo‐managed rhinos

Assessing zoo giraffe survivorship: Methodological aspects, historical improvement and a rapid demographic shift

Giraffe have been kept in zoos for a long time. They have traditionally been considered difficult to maintain due to various husbandry requirements, including their nature as intrinsic browsers. However, zoo animals are expected to achieve higher survivorship than free-ranging conspecifics due to protection against dangers that would be experienced in their natural habitat. Global zoo giraffe data was analysed for historical developments of juvenile and adult survivorship, assessing the data with various demographic measures and comparing it to that of populations from natural habitats. Additionally, zoo population structure was analysed, in particular with respect to two events that occurred in parallel in 2014—a recommendation to restrict the number of new offspring given by the European Endangered Species Programme (EEP) studbook coordinator and the culling of a designated ‘surplus’ giraffe at Copenhagen Zoo that attracted global media attention. Both juvenile and adult giraffe survivorship has increased over time, suggesting advances in giraffe husbandry. For juveniles, this process has been continuous, whereas for adults the major progress has been in the most recent cohort (from 2000 onwards), in parallel with the publication of various husbandry guidelines. Zoo giraffe survivorship is now generally above that observed in natural habitats. Simple survivorship analyses appear suitable to describe these developments. Since 2014, the global giraffe population has undergone a rapid demographic shift from a growing to an ageing population, indicating a drastic limitation of reproduction rather than a system where reproduction is allowed and selected animals are killed (and possibly fed to carnivores). Thus, giraffe are both a showcase example for the historical progress made in zoo animal husbandry due to efforts of the zoo community and serve as an example to discuss implications of different methods of zoo population management

The historical development of zoo elephant survivorship

In the discussion about zoo elephant husbandry, the report of Clubb et al. (2008, Science 322: 1649) that zoo elephants had a “compromised survivorship” compared to certain non-zoo populations is a grave argument, and was possibly one of the triggers of a large variety of investigations into zoo elephant welfare, and changes in zoo elephant management. A side observation of that report was that whereas survivorship in African elephants (Loxodonta africana) improved since 1960, this was not the case in Asian elephants (Elephas maximus). We used historical data (based on the Species360 database) to revisit this aspect, including recent developments since 2008. Assessing the North American and European populations from 1910 until today, there were significant improvements of adult (≥10 years) survivorship in both species. For the period from 1960 until today, survivorship improvement was significant for African elephants and close to a significant improvement in Asian elephants; Asian elephants generally had a higher survivorship than Africans. Juvenile (<10 years) survivorship did not change significantly since 1960 and was higher in African elephants, most likely due to the effect of elephant herpes virus on Asian elephants. Current zoo elephant survivorship is higher than some, and lower than some other non-zoo populations. We discuss that in our view, the shape of the survivorship curve, and its change over time, are more relevant than comparisons with specific populations. Zoo elephant survivorship should be monitored continuously, and the expectation of a continuous trend towards improvement should be met

Comparative analyses of longevity and senescence reveal variable survival benefits of living in zoos across mammals

While it is commonly believed that animals live longer in zoos than in the wild, this assumption has rarely been tested. We compared four survival metrics (longevity, baseline mortality, onset of senescence and rate of senescence) between both sexes of free-ranging and zoo populations of more than 50 mammal species. We found that mammals from zoo populations generally lived longer than their wild counterparts (84% of species). The effect was most notable in species with a faster pace of life (i.e. a short life span, high reproductive rate and high mortality in the wild) because zoos evidently offer protection against a number of relevant conditions like predation, intraspecific competition and diseases. Species with a slower pace of life (i.e. a long life span, low reproduction rate and low mortality in the wild) benefit less from captivity in terms of longevity; in such species, there is probably less potential for a reduction in mortality. These findings provide a first general explanation about the different magnitude of zoo environment benefits among mammalian species, and thereby highlight the effort that is needed to improve captive conditions for slow-living species that are particularly susceptible to extinction in the wild

Reproductive seasonality in primates: patterns, concepts and unsolved questions

Primates, like other mammals, exhibit an annual reproductive pattern that ranges from strictly seasonal breeding to giving birth in all months of the year, but factors mediating this variation are not fully understood. We applied both a categorical description and quantitative measures of the birth peak breadth based on daily observations in zoos to characterise reproductive seasonality in 141 primate species with an average of 941 birth events per species. Absolute day length at the beginning of the mating season in seasonally reproducing species was not correlated between populations from natural habitats and zoos. The mid‐point of latitudinal range was a major factor associated with reproductive seasonality, indicating a correlation with photoperiod. Gestation length, annual mean temperature, natural diet and Malagasy origin were other important factors associated with reproductive seasonality. Birth seasons were shorter with increasing latitude of geographical origin, corresponding to the decreasing length of the favourable season. Species with longer gestation periods were less seasonal than species with shorter ones, possibly because shorter gestation periods more easily facilitate the synchronisation of reproductive activity with annual cycles. Habitat conditions with higher mean annual temperature were also linked to less‐seasonal reproduction, independently of the latitude effect. Species with a high percentage of leaves in their natural diet were generally non‐seasonal, potentially because the availability of mature leaves is comparatively independent of seasons. Malagasy primates were more seasonal in their births than species from other regions. This might be due to the low resting metabolism of Malagasy primates, the comparatively high degree of temporal predictability of Malagasy ecosystems, or historical constraints peculiar to Malagasy primates. Latitudinal range showed a weaker but also significant association with reproductive seasonality. Amongst species with seasonal reproduction in their natural habitats, smaller primate species were more likely than larger species to shift to non‐seasonal breeding in captivity. The percentage of species that changed their breeding pattern in zoos was higher in primates (30%) than in previous studies on Carnivora and Ruminantia (13 and 10%, respectively), reflecting a higher concentration of primate species in the tropics. When comparing only species that showed seasonal reproduction in natural habitats at absolute latitudes ≤11.75°, primates did not differ significantly from these two other taxa in the proportion of species that changed to a less‐seasonal pattern in zoos. However, in this latitude range, natural populations of primates and Carnivora had a significantly higher proportion of seasonally reproducing species than Ruminantia, suggesting that in spite of their generally more flexible diets, both primates and Carnivora are more exposed to resource fluctuation than ruminants