COMPARISON OF VERTICAL AERIAL PHOTOGRAPHIC AND GROUND CENSUSES OF STELLER SEA LIONS AT AÑO NUEVO ISLAND, JULY 1990-1993

Counts of Steller sea lion ( Eumetopias jubatus ) pups and non-pups (adults and juveniles) from aerial photographs of rookeries at Año Nuevo Island between 1990 and 1993 were significantly higher than those made on the ground. Based on regression of natural logs of photographic counts versus year, the number of pups declined at a rate of −0.099yr while non-pup numbers declined at −0.315/yr. Examination of ground count data for the same period revealed a significant decline in non-pups (−0.139/yr), but no trend was detected in the ground counts of pups. The regression coefficients from photographic and ground counts of non-pups did not differ significantly. Power analyses using the program TRENDS indicated that detectable rates of change in abundance from four annual surveys were much lower for counts of pups than counts of non-pups where sampling precision was based on fits to linear models.     

Captive killer whale (Orcinus orca) survival

Killer whales (Orcinus orca) were first placed into captivity in 1961 and are now found in theme parks around the world. Despite successful breeding of captive killer whales since 1985 there is growing concern for their welfare in captivity, which often includes claims of poor survival. We employed Kaplan‐Meier and Cox Proportional hazards models and annual survival rate analyses on 201 captive killer whales to discern how sex, facility (U.S. vs. foreign), captive‐born vs. wild‐captured, pre‐ vs. post‐1 January 1985, and animal age upon entering captivity affect survival. Overall median survival estimate was 6.1 yr, with no difference between male and female survival. Killer whales in U.S. facilities (12.0 yr) demonstrated a significantly higher median survival than those in foreign facilities (4.4 yr), as did whales entering captivity post‐1 January 1985 (11.8 yr) vs. those entering prior to 1 January 1985 (3.9 yr). Median survival for captive‐born (14.1 yr) was significantly higher than wild‐captured killer whales (5.5 yr), though the two failed to differ among the post‐1 January 1985 cohort. Facility location and pre‐ vs. post‐1 January 1985 were predictors of the hazard rate. Survival of captive killer whale cohorts has generally improved through time, although survival to age milestones are poor when compared to wild killer whales.     

ACCLIMATION TO CAPTIVITY: A QUANTITATIVE ESTIMATE BASED ON SURVIVAL OF BOTTLENOSE DOLPHINS AND CALIFORNIA SEA LIONS

An estimate of how long marine mammals need to acclimate to captivity would permit more precise comparisons of husbandry practices, yet no quantitative analysis of acclimation has been performed. Therefore, we estimated the duration of acclimation to captivity for bottlenose dolphins (BD) and California sea lions (CSL) by comparing 5-d survival rates during the first 90 d of captivity with a survival rate based on days 91-365 in captivity. Wild-born BD (n = 1,270) and CSL (n = 1,650) acclimate to captivity in approximately 35 and 40 d, respectively, whereas captive born BD (n = 332) and CSL (n = 992) acclimate in approximately 50 and 40 d, respectively. When transferred between two institutions, BD (n = 911) acclimated in the same amount of time (45 d) as when first transferred from the wild, whereas transferred CSL (n = 336) acclimated more rapidly (15 VJ. 40 d) than when first transferred from the wild. Based on results from these two species, a 60-d acclimation period is recognized as a distinct interval of relatively high mortality that should be treated separately from long-term survival estimates when evaluating husbandry practices of ocean-aria and zoos.     

Comparative Skull Analysis Suggests Species-Specific Captivity-Related Malformation in Lions (Panthera leo)

Lion (Panthera leo) populations have dramatically decreased worldwide with a surviving population estimated at 32,000 across the African savannah. Lions have been kept in captivity for centuries and, although they reproduce well, high rates of stillbirths as well as morbidity and mortality of neonate and young lions are reported. Many of these cases are associated with bone malformations, including foramen magnum (FM) stenosis and thickened tentorium cerebelli. The precise causes of these malformations and whether they are unique to captive lions remain unclear. To test whether captivity is associated with FM stenosis, we evaluated 575 lion skulls of wild (N = 512) and captive (N = 63) origin. Tiger skulls (N = 276; 56 captive, 220 wild) were measured for comparison. While no differences were found between males and females or between subadults and adults in FM height (FMH), FMH of captive lions (17.36±3.20 mm) was significantly smaller and with greater variability when compared to that in wild lions (19.77±2.11 mm). There was no difference between wild (18.47±1.26 mm) and captive (18.56±1.64 mm) tigers in FMH. Birth origin (wild vs. captive) as a factor for FMH remained significant in lions even after controlling for age and sex. Whereas only 20/473 wild lions (4.2%) had FMH equal to or smaller than the 5^th percentile of the wild population (16.60 mm), this was evident in 40.4% (23/57) of captive lion skulls. Similar comparison for tigers found no differences between the captive and wild populations. Lions with FMH equal to or smaller than the 5^th percentile had wider skulls with smaller cranial volume. Cranial volume remained smaller in both male and female captive lions when controlled for skull size. These findings suggest species- and captivity-related predisposition for the pathology in lions.     

Characterization of Ovarian Steroid Patterns in Female African Lions (Panthera leo), and the Effects of Contraception on Reproductive Function

Because of poor reproduction after the lifting of an 8-year breeding moratorium, a biomedical survey of female lions in U.S. zoos was initiated in 2007. Fecal estrogen (FEM), progestagen (FPM) and glucocorticoid (FGM) metabolites were analyzed in samples collected 3–4 times per wk from 28 lions at 17 facilities (0.9–13.8 yr of age) for 4 mo—3.5 yr and body weights were obtained ~monthly from 17 animals at eight facilities (0.0–3.0 yr of age). Based on FEM, estrous cycle length averaged 17.5 ± 0.4 d in duration, with estrus lasting 4.4 ± 0.2 d. All but one female exhibited waves of estrogenic activity indicative of follicular activity; however, not all females expressed estrous behaviors (73%), suggesting silent estrus was common. Female lions experienced puberty earlier than expected; waves of estrogenic activity were observed as young as 1.1 yr of age, which may be related to a faster growth rate of captive vs. wild lions. Mean gestation length was 109.5 ± 1.0 d, whereas the non-pregnant luteal phase was less than half (46.0 ± 1.2 d). Non-mating induced increases in FPM were observed in 33% of females housed without a male, consistent with spontaneous ovulation. A number of study animals had been contracepted, and the return to cyclicity after treatment withdrawal, while variable, was ~4.0 yr and longer than the 1-yr expected efficacy, especially for those implanted with Suprelorin. For FGM, there were no differences in overall, baseline or peak mean concentrations among the age groups or across seasons, nor were there any relationships between reproductive parameters and FGM concentrations. Overall, results suggest that poor reproduction in lions after the breeding moratorium was not related to altered adrenal or ovarian steroid activity, but for some females may have been a consequence of individual institutions’ management decisions.     

Swimming speed, respiration rate, and estimated cost of transport in adult killer whales

The physiology of free-ranging cetaceans is difficult to study and as a consequence, data on the energetics of these animals are limited. To better understand the energetic cost of swimming in killer whales, total cost of transport ( COT ) was estimated from swimming speeds and respiration rates from wild adult northern resident killer whales ( Orcinus orca ) and reported values of oxygen consumption in captive whales. Respiration rate (breaths per minute) was positively correlated with swimming speed (meters per second), while mass-specific COT (Joules per kilogram per meter) decreased with speed. Lack of data on very fast-swimming animals hindered assessment of the exact speed at which COT was minimal. However, minimum mass-specific COT for killer whales in the present study approached those predicted by a previously published allometric equation for marine mammals, and corresponded to "optimal" swimming speeds of 2.6–3 m/s. Interestingly, the observed average swimming speed (1.6 m/s) was lower than predicted optimal swimming speed. Finally, females with dependent calves had higher respiration rates than females without calves. These findings could be due to synchronous breathing with calves or could result from increased costs of lactation and swimming with a calf in echelon formation. Consequently, females with calves may have much greater COT at optimal swimming speeds than females without calves.     

Nutritional and behavioral effects of gorge and fast feeding in captive lions

Nonhuman animals in captivity manifest behaviors and physiological conditions that are not common in the wild. Lions in captivity face problems of obesity, inactivity, and stereotypy. To mediate common problems of captive lions, this study implemented a gorge and fast feeding schedule that better models naturalistic patterns: African lions (Panthera leo) gradually adapted from a conventional feeding program to a random gorge and fast feeding schedule. Digestibility increased significantly and food intake and metabolizable energy intake correspondingly decreased. Lions also showed an increase in appetitive active behaviors, no increase in agonistic behavior, and paced half as frequently on fast days as on feeding days. Thus, switching captive lions to a gorge and fast feeding schedule resulted in improved nutritional status and increased activity.     

RANGE-WIDE SURVEY AND ESTIMATION OF TOTAL NUMBER OF STELLER SEA LIONS IN 1989

Abstract: The first range-wide survey of Steller (northern) sea lions ( Eumetopias jubatus ) was completed in 1989 with a total of 68,094 adult and juvenile (nonpup) Steller sea lions counted. This total count includes 10,000 in Russia (15% of the range-wide count), 47,960 in Alaska (70%), 6,109 in British Columbia (9%), 2,261 in Oregon (3%), and 1,764 in California (3%). A range-wide pup count was not obtained. We estimated the 1989 world population based on a calculation for total pups and obtained a range-wide estimate of 116,000 total animals, or about 39–48% of the 240,000-300,000 estimated 30 yr ago.     

How long do dolphins live? Survival rates and life expectancies for bottlenose dolphins in zoological facilities vs. wild populations

Survival rates and life expectancies are commonly agreed upon indicators of well‐being for animals in zoological facilities, but even the most recent survival statistics for bottlenose dolphins (Tursiops truncatus) in marine mammal parks and aquariums use data that are now more than 25 yr old. The current study provides a comprehensive assessment of life expectancy and survival rates for bottlenose dolphins in U.S. zoological facilities from 1974 to 2012, utilizing three different analyses (annual survival rate, age‐at‐death, and Kaplan‐Meier), examining historical trends, and comparing to comparable data from wild populations. Both survival rate and life expectancy for dolphins in zoological facilities increased significantly over the past few decades, with a modern ASR of 0.972, and mean and median life expectancies calculated via Kaplan‐Meier of 28.2 and 29.2 yr, respectively. Survival rates and life expectancies for dolphins in U.S. zoological facilities today are at least as high as those for the wild dolphin populations for which there are comparable data.     

SURVIVORSHIP PATTERNS IN THREE SPECIES OF CAPTIVE CETACEANS

Survival rates for three species of captive cetaceans are reported, based on records of dates of capture, birth, and death of individual animals. The annual survival rate was 0.93 for bottlenose dolphins and killer whales and 0.94 for white whales. Confidence limits of these estimates are discussed. Differences in survival rates between institutions were significant for bottlenose dolphins only. Calf survival for bottlenose dolphins was lower than non-calf survival. Survivorship of male killer whales was significantly less than survivorship of female killer whales; sex-specific survival rates were similar for the other two species. Estimates of average or maximum longevity alone were not useful in comparing rates of survival. Because survival in the first year of captivity may be lower than subsequent years, estimates of the expected lifespan, based on data from the first few years of captivity, may be biased.     

Temporal variability in organochlorine contamination in blood and blubber of captive Steller sea lions (Eumetopias jubatus)

Three adult captive Steller sea lions (SSL; two females, one male) housed in Alaska were longitudinally sampled for organochlorine contaminant (OC) analysis in both blubber (n= 19) and blood (n= 69) over a 2 yr period (March 2001 to March 2003). Blood OC concentrations were similar between individuals, and exhibited significant increases in summer months (July through September) relative to winter (January through March). Additionally, paired blood and blubber sample (n= 18) OC were significantly related for all animals. The relationship of blubber OC concentrations to lipid content was significant in all animals. Although limited to a small number of animals, our study results indicate that in SSLs, blood OC were both consistent among all animals and likely changed in association with physiologically driven metabolism of blubber.     

A genetically distinct lion (Panthera leo) population from Ethiopia

Lion (Panthera leo) numbers are in serious decline and two of only a handful of evolutionary significant units have already become extinct in the wild. However, there is continued debate about the genetic distinctiveness of different lion populations, a discussion delaying the initiation of conservation actions for endangered populations. Some lions from Ethiopia are phenotypically distinct from other extant lions in that the males possess an extensive dark mane. In this study, we investigated the microsatellite variation over ten loci in 15 lions from Addis Ababa Zoo in Ethiopia. A comparison with six wild lion populations identifies the Addis Ababa lions as being not only phenotypically but also genetically distinct from other lions. In addition, a comparison of the mitochondrial cytochrome b (CytB) gene sequence of these lions to sequences of wild lions of different origins supports the notion of their genetic uniqueness. Our examination of the genetic diversity of this captive lion population shows little effect of inbreeding. Immediate conservation actions, including a captive breeding programme designed to conserve genetic diversity and maintain the lineage, are urgently needed to preserve this unique lion population.     

Accuracy and precision of age determination using growth layer groups for California sea lions (Zalophus californianus) with known ages

Age determination from counts of growth layer groups (GLGs) in tooth dentine is a common method for aging marine mammals. Using known‐aged animals, we validated this method for acid etched teeth of California sea lions (CSLs), Zalophus californianus. Between 1991 and 2013, the upper left canine (n = 33) was collected opportunistically during necropsy from animals tagged or branded as pups that later died. Overall, 55%–61% of age estimates by GLG counting were within 1 yr of the known‐age in the sample of 1–30‐yr‐old CSLs. Accuracy of age estimates was found to be dependent on age of the CSLs, however. 71%–79% of age estimates were within 1 yr of the known‐age in CSLs <10 yr old. These findings support the validity of counting GLGs to estimate age for CSLs <10 yr old to within 1 yr of accuracy.     

ENVIRONMENTAL FACTORS INFLUENCING THE DISTRIBUTION OF SOUTHERN RIGHT WHALES (EUBALAENA AUSTRALIS) ON THE SOUTH COAST OF SOUTH AFRICA I: BROAD SCALE PATTERNS

Aerial surveys over the last 32 yr have shown that the distribution of southern right whales Eubalaena australis along the south coast of South Africa is markedly discontinuous, but highly predictable. A GIS was used at a variety of scales to investigate whether this pattern was related to environmental characteristics. Whale distribution was analyzed as density per 20-min bin of longitude over two temporal and spatial scales, namely 15 bins for 32 yr, and a wider scale but shorter time period, 23 bins for 19 yr, as well as using three years of GPS accuracy data (15 bins) for finer scale analysis. Environmental factors tested were depth, distance from shore, sea floor slope, protection from swell, protection from wind, and shore type. The majority of whales were concentrated in areas that provided reasonable protection from open ocean swell and seasonal winds, and had sedimentary floors with gentle slopes. They generally avoided exposed rocky shorelines. Cow-calf pairs were found significantly closer to shore and in shallower water than unaccompanied whales, particularly off sandy beaches. Habitat choice at this time of year may be related both to energy conservation for calves and lactating females (calm sea conditions) and to protection of the new-born.     

Interactions between killer whales (<Emphasis Type="Italic">Orcinus orca</Emphasis>) and Steller sea lions (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>) in the vicinity of Brat Chirpoev Island,Kuril Islands

The behaviors of breeding Steller sea lions in response to encounters with killer whales near the shore were observed on Brat Chirpoev Island, Kuril Islands between May and July 2002–2007. Approaches by killer whales and sea lion behavior was observed visually and recorded. Killer whales approached the rookery 104 times during the entire period of observations (289 days). In most cases (n = 95), beached sea lions did not show any apparent reactions to the presence of killer whales, and there were no observed interactions. Sea lions showed agitation during nine of the approaches; five of these events were considered to be predation attempts. The killer whales attacked the sea lions three times, however all the attacks were unsuccessful. We recorded two different types of responses towards the killer whales: (1) beaching on the shore (three times) and (2) mass exodus from the rookery with subsequent formation of a tight, actively swimming and vocalizing group (six times). The latter is the first recorded observation of this behavior for Steller sea lions. The observation suggests a low degree of interactions between these two species near the studied rookery. Despite the numerous observations of killer whales near the rookery, there were no observations of direct predation on sea lions. It is likely the killer whale predation has little or no direct impact on the Steller sea lion population on Brat Chirpoev Islands during the breeding period.     

FORAGING STRATEGIES OF SYMPATRIC KILLER WHALE (ORCINUS ORCA) POPULATIONS IN PRINCE WILLIAM SOUND, ALASKA

Killer whales ( Orcinus orca ) feed on a wide variety of fish, cephalopods, and marine mammals throughout their cosmopolitan range; however, the dietary breadth that characterizes the species is not reflected in all populations. Here, we present the findings of a 14-yr study of the diet and feeding habits of killer whales in Prince William Sound, Alaska. Two non-associating forms of killer whale, termed resident and transient (Bigg et al. 1987), were identified. All prey seen taken by transients were marine mammals, including harbor seals ( Phoca vitulina ), Dall's porpoises ( Phocoenoides dalli ), Steller sea lions ( Eumetopias jubatus ), and harbor porpoises ( Phocoena phocoena ). Resident killer whales appeared to prey principally on salmon ( Oncorhynchus spp.), preferring coho salmon ( O. kisutch ) over other, more abundant salmon species. Pacific herring ( Clupea pallasi ) and Pacific halibut ( Hippocampus stenolepis ) were also taken. Resident killer whales frequently were seen to interact in non-predatory ways with Steller sea lions and Dall's porpoises, while transients were not. Differences in the social organization and behavior of the resident and transient killer whales in Prince William Sound are discussed in the light of the dietary differences documented here.     

SEXUAL COMPOSITION and ANALYSIS OF REPRODUCTIVE FEMALES IN THE NORTH ATLANTIC RIGHT WHALE, EUBALAENA GLACIALIS, POPULATION

To test hypotheses involving reproduction and demographics, the sex of individuals must be established, but many species of Cetacea are not obviously dimorphic. In the North Atlantic right whale, Eubalaena glacialis , population, the sex of 61 males and 55 females had been determined previously by observation of the urogenital region, and the sex of 43 more females had been inferred from repeated sightings with a calf. To confirm the sex of some of these animals and to identify the sex of mote animals, genomic DNA was isolated from skin samples of 95 individual right whales (54 from among those described above and 41 additional recognizable individuals). The DNA was surveyed using the human Y-chromosome probe pDP1007. With Eco RI-digested DNA, a clear, sex-discriminating banding pattern was apparent. This method verified the sex of all 54 animals whose sex was previously known or inferred and identified the sex of an additional 41 recognizable individuals. A total of 89 male and 111 female right whales was identified in the population. The most unbiased estimate of sex ratio available is the 36 male and 34 female calves identified by genital morphology and DNA techniques. The sex ratio of this sample does not differ significantly from unity (P = 0.811). Only 38% (58/152) of the females in the North Atlantic population are known to have been reproductively successful compared with 54% in the population of right whales in the western South Atlantic. The population growth rate reported for the North Atlantic population is only 33% of that reported for right whales in the South Atlantic. Thirteen adult North Atlantic females have been identified that have not been known to calve during the past 11 yr. These data suggest that the absence of measurable recovery may be due to a combination of fewer actively reproducing females and lower reproductive rates of some females.     

STELLER SEA LION STATUS AND TREND IN SOUTHEAST ALASKA: 1979–1997

Steller sea lion (Eumetopias jubatus) numbers in the United States declined by about 75% over the past 20+ yr. They are classified, under the U. S. Endangered Species Act, as “threatened” in the eastern portion of their range and as “endangered” in the western portion. We analyzed trends in numbers of pup and non-pup Steller sea lions counted in Southeast Alaska between 1979 and 1997. Sea lion numbers, based on counts of pups on rookeries, increased by an average of 5.9% per year between 1979 and 1997. However, numbers of pups increased at a much slower rate (+ 1.7% per year) between 1989 and 1997. For counts of non-pup Steller sea lions we used models that controlled for the effects of date, time, and tide at the time of the survey to analyze trends. This technique reduced bias and increased precision of the resulting trend estimates. Numbers of sea lions were stable (+0.5%) between 1989 and 1996, based on counts of non-pups. We estimated the Southeast Alaska breeding population of Steller sea lions at about 19,000 animals of all ages in 1997, a level that is probably near the highest in recorded history.     

African Wild Dogs as a Fugitive Species: Playback Experiments Investigate How Wild Dogs Respond to their Major Competitors

It has been suggested that African wild dogs Lycaon pictus need exceptionally large home ranges (and hence occur at such low densities) because they are limited by competition with larger sympatric carnivores, namely lions Panthera leo and spotted hyenas Crocuta crocuta. To investigate this relationship at a proximate level and explore which factors mediate it, we conducted audio playback experiments examining how wild dogs responded to the simulated proximity of either lions or hyenas. The principle finding was that wild dogs consistently moved directly away from lion roars, but when played hyena whoops either stood their ground or, later, moved off in a random direction. These results suggest that lions represent an immediate high‐level threat to wild dogs that is invariably best avoided, whilst the threat from hyenas may not be so great or perhaps is simply unavoidable. Wild dogs appeared to make some assessment of ambush risk during interactions with lions, illustrated by the varying latency to their retreat in habitats of differing vegetation density (and hence ambush potential). Additionally, packs with younger pups were more likely to alarm call and exhibited a slower rate of retreat in the hour following exposure to lion roars. Other variables investigated (competitor group size, lion sex, presence of pups) failed to explain variation in wild dogs’ responses.     

The Three-Dimensional Morphological Effects of Captivity

Many captive animals are fed diets that are drastically different in mechanical properties than their wild diet. Most captive pantherines are fed a nutritionally supplemented diet consisting almost entirely of ground meat. While many zoos supplement this diet with bones, the fact remains that large captive felids are fed diets that require substantially less masticatory effort than those of their wild counterparts. The osteological effects of this dietary difference have not been fully evaluated. To this end, we compared linear measurements and 3D geometric morphometric landmarks of captive and wild lions and tigers. Using Principal Component (PC) analysis of the linear measurements, not only were the sexes and species statistically distinct, but so too was the population clearly divisible in terms of captivity status. The 3D analysis supported these findings: although the most influential variable in the sample (PC1, 21.5% of the variation) separates the two species, the second most influential contributor (PC2) to the overall skull shape is driven not by the sex differences in these highly dimorphic species, but rather by their captivity status. In fact, captivity status drives nearly twice as much of the 3D variation as sexual dimorphism (14.8% vs. 8.0% for PC2 vs. PC3). Thus the shape is influenced nearly twice as much by whether the animal was captive or wild than by whether it was male or female. If a causal relationship can be demonstrated between dietary mechanical properties and morphology, people who oversee the diets of captive carnivores should consider modifying these diets to account for not only nutritional but also the mechanical properties of a carcass-based diet as well. In addition to the husbandry implications, our analyses show the ways in which captive specimens are different than their wild counterparts – findings that have implications for morphologists when considering anatomical samples.