High Natality Rates of Endangered Steller Sea Lions in Kenai Fjords,Alaska and Perceptions of Population Status in the Gulf of Alaska

Steller sea lions experienced a dramatic population collapse of more than 80% in the late 1970s through the 1990s across their western range in Alaska. One of several competing hypotheses about the cause holds that reduced female reproductive rates (natality) substantively contributed to the decline and continue to limit recovery in the Gulf of Alaska despite the fact that there have been very few attempts to directly measure natality in this species. We conducted a longitudinal study of natality among individual Steller sea lions (n = 151) at a rookery and nearby haulouts in Kenai Fjords, Gulf of Alaska during 2003–2009. Multi-state models were built and tested in Program MARK to estimate survival, resighting, and state transition probabilities dependent on whether or not a female gave birth in the previous year. The models that most closely fit the data suggested that females which gave birth had a higher probability of surviving and giving birth in the following year compared to females that did not give birth, indicating some females are more fit than others. Natality, estimated at 69%, was similar to natality for Steller sea lions in the Gulf of Alaska prior to their decline (67%) and much greater than the published estimate for the 2000s (43%) which was hypothesized from an inferential population dynamic model. Reasons for the disparity are discussed, and could be resolved by additional longitudinal estimates of natality at this and other rookeries over changing ocean climate regimes. Such estimates would provide an appropriate assessment of a key parameter of population dynamics in this endangered species which has heretofore been lacking. Without support for depressed natality as the explanation for a lack of recovery of Steller sea lions in the Gulf of Alaska, alternative hypotheses must be more seriously considered.     

Investigating life‐history traits of Steller sea lions with multistate hidden Markov mark–recapture models: Age at weaning and body size effects

The duration of offspring care is critical to female fitness and population resilience by allowing flexibility in life‐history strategies in a variable environment. Yet, for many mammals capable of extended periods of maternal care, estimates of the duration of offspring dependency are not available and the relative importance of flexibility of this trait on fitness and population viability has rarely been examined. We used data from 4,447 Steller sea lions Eumetopias jubatus from the Gulf of Alaska and multistate hidden Markov mark–recapture models to estimate age‐specific weaning probabilities. Maternal care beyond age 1 was common: Weaning was later for animals from Southeast Alaska (SEAK) and Prince William Sound (PWS, weaning probabilities: 0.536–0.648/0.784–0.873 by age 1/2) compared with animals born to the west (0.714–0.855/0.798–0.938). SEAK/PWS animals were also smaller than those born farther west, suggesting a possible link. Females weaned slightly earlier (+0.080 at age 1 and 2) compared with males in SEAK only. Poor survival for weaned versus unweaned yearlings occurred in southern SEAK (female survival probabilities: 0.609 vs. 0.792) and the central Gulf (0.667 vs. 0.901), suggesting poor conditions for juveniles in these areas. First‐year survival increased with neonatal body mass (NBM) linearly in the Gulf and nonlinearly in SEAK. The probability of weaning at age 1 increased linearly with NBM for SEAK animals only. Rookeries where juveniles weaned at earlier ages had lower adult female survival, but age at weaning was unrelated to population trends. Our results suggest the time to weaning may be optimized for different habitats based on long‐term average conditions (e.g., prey dynamics), that may also shape body size, with limited short‐term plasticity. An apparent trade‐off of adult survival in favor of juvenile survival and large offspring size in the endangered Gulf of Alaska population requires further study.     

DISPERSAL, ROOKERY FIDELITY, AND METAPOPULATION STRUCTURE OF STELLER SEA LIONS (EUMETOPIAS JUBATUS) IN AN INCREASING AND A DECREASING POPULATION IN ALASKA

Over the past 24 yr, 8,596 Steller sea lion ( Eumetopias jubatus ) pups were branded on their natal rookeries throughout Alaska with the objectives of determining survival rates, recruitment, movements, and site fidelity. Our objectives here were to examine the extent of dispersal of Steller sea lions away from their natal rookeries, movements between stocks, and degree of natal rookery fidelity. Pups (<1 yr old) usually remained within 500 km of their natal rookery. Branded juveniles dispersed widely and were resighted at distances up to 1,785 km from their natal rookeries. Adults generally remained within 500 km of their natal rookeries. No interchange of breeding animals between the ES (eastern stock) and WS (western stock) was observed. Although natal rookery fidelity was prevalent, 33% of the 12 observations of females branded in the WS during 1987–1988 and 19% of the 29 observations of females branded in the ES during 1994–1995 were observed with newly born pups at sites other than their natal rookeries. Steller sea lions generally conformed to the metapopulation concept as depicted by Hanski and Simberloff (1997), with local breeding populations (rookeries) and movements among these local populations having the potential of affecting local dynamics.     

Age Specific Survival Rates of Steller Sea Lions at Rookeries with Divergent Population Trends in the Russian Far East

After a dramatic population decline, Steller sea lions have begun to recover throughout most of their range. However, Steller sea lions in the Western Aleutians and Commander Islands are continuing to decline. Comparing survival rates between regions with different population trends may provide insights into the factors driving the dynamics, but published data on vital rates have been extremely scarce, especially in regions where the populations are still declining. Fortunately, an unprecedented dataset of marked Steller sea lions at rookeries in the Russian Far East is available, allowing us to determine age and sex specific survival in sea lions up to 22 years old. We focused on survival rates in three areas in the Russian range with differing population trends: the Commander Islands (Medny Island rookery), Eastern Kamchatka (Kozlov Cape rookery) and the Kuril Islands (four rookeries). Survival rates differed between these three regions, though not necessarily as predicted by population trends. Pup survival was higher where the populations were declining (Medny Island) or not recovering (Kozlov Cape) than in all Kuril Island rookeries. The lowest adult (> 3 years old) female survival was found on Medny Island and this may be responsible for the continued population decline there. However, the highest adult survival was found at Kozlov Cape, not in the Kuril Islands where the population is increasing, so we suggest that differences in birth rates might be an important driver of these divergent population trends. High pup survival on the Commander Islands and Kamchatka Coast may be a consequence of less frequent (e.g. biennial) reproduction there, which may permit females that skip birth years to invest more in their offspring, leading to higher pup survival, but this hypothesis awaits measurement of birth rates in these areas.     

SEASONAL DISTRIBUTION OF STELLER'S SEA LIONS AT ROOKERIES AND HAUL-OUT SITES IN ALASKA

The Steller's sea lion population has declined by 60%-70% over much of Alaska since the late 1970s. Overlap in species composition and sizes of fishes consumed by sea lions and harvested by commercial fisheries, particularly during winter, has led to examination of potential interaction between commercial fisheries and Steller's sea lions. Abundance and distribution data for Steller's sea lions in Alaska were derived from aerial surveys conducted during the breeding season, mid-June to early July 1992, 1994, and 1996. To study winter distribution of sea lions, we conducted aerial surveys during March 1993, November-December 1994, and March 1999. We counted about one-half as many sea lions during winter surveys compared to the breeding-season surveys. Numbers of sea lions at rookery sites dropped off considerably during winter, whereas numbers at haul-out sites did not. We found little evidence of large-scale, seasonal movement, at least for the western stock of sea lions. Rather, differences between summer and winter distribution were primarily a function of sea lions dispersing to local haul-out sites during the winter. Terrestrial sites, both rookeries and haul-outs, clearly are important to Steller's sea lions during the entire year. Individual sites may be occupied year-round or only during particular times of year.     

Assessment of Competition between Fisheries and Steller Sea Lions in Alaska Based on Estimated Prey Biomass,Fisheries Removals and Predator Foraging Behaviour

A leading hypothesis to explain the dramatic decline of Steller sea lions (Eumetopias jubatus) in western Alaska during the latter part of the 20th century is a change in prey availability due to commercial fisheries. We tested this hypothesis by exploring the relationships between sea lion population trends, fishery catches, and the prey biomass accessible to sea lions around 33 rookeries between 2000 and 2008. We focused on three commercially important species that have dominated the sea lion diet during the population decline: walleye pollock, Pacific cod and Atka mackerel. We estimated available prey biomass by removing fishery catches from predicted prey biomass distributions in the Aleutian Islands, Bering Sea and Gulf of Alaska; and modelled the likelihood of sea lions foraging at different distances from rookeries (accessibility) using satellite telemetry locations of tracked animals. We combined this accessibility model with the prey distributions to estimate the prey biomass accessible to sea lions by rookery. For each rookery, we compared sea lion population change to accessible prey biomass. Of 304 comparisons, we found 3 statistically significant relationships, all suggesting that sea lion populations increased with increasing prey accessibility. Given that the majority of comparisons showed no significant effect, it seems unlikely that the availability of pollock, cod or Atka mackerel was limiting sea lion populations in the 2000s.     

Adult Steller sea lion mortality on rookeries in the Russian Far East, 2002–2010

We used visual observations during nine field seasons to determine the causes and death rate of Steller sea lions older than 1 year on eight rookeries in the Russian Far East. The average annual death rate was 0.48% of the total recorded Steller sea lions on shore and varied among rookeriesy. The mortality did not exceed 0.29% in six rookeries, but reached 0.80–1.19% in two rookeries. Individuals of all age and sex groups were recorded among dead animals. Bulls died as a result of territorial conflicts, and cows and young lions were crushed or suffocated by bulls during copulation. The death of nearly half of the females was attributed to 11 bulls.     

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.     

DEVELOPMENT OF DISPERSAL, MOVEMENT PATTERNS, AND HAUL-OUT USE BY PUP AND JUVENILE STELLER SEA LIONS (EUMETOPIAS JUBATUS) IN ALASKA

Population declines of Steller sea lions ( Eumetopias juhatus ) in western Alaska (west of 144°W) may be a result of reduced juvenile survival. We used satellite telemetry to study the at-sea distribution and movement patterns of pup (1.6–11.9 mo) and juvenile (12.0–35.1 mo) Steller sea lions. We studied trip distance, duration, and interhaul-out movements of sea lions in relation to age, sex, and month of year in the decreasing western population (WP; Prince William Sound, Kodiak, Aleutian Islands, Alaska) and the increasing eastern population (EP; Southeast Alaska). We deployed 103 satellite transmitters (29 WP; 74 EP) on sea lions between 1998 and 2001. Round trip distance and duration increased with age, trip distance was greater in the WP than the EP, trip duration was greater for females than males, and haul-out use was clustered. Changes in round trip distance and duration occurred from April to June for all age classes studied indicating that the annual timing of weaning may be less variable than the age of weaning. Overall, 90% of round trips were ≤ 15 km from haul-outs and 84% were <20 h, indicating nearshore areas adjacent to haulouts are critical to the developing juvenile.     

COUNTING STELLER SEA LION PUPS IN ALASKA: AN EVALUATION OF MEDIUMFORMAT, COLOR AERIAL PHOTOGRAPHY

Estimates of Steller sea lion ( Eumetopias jubatus ) pup production are valuable for estimating population trend and size. Currently in Alaska, pups are counted by visiting rookeries, driving older animals into the water, then walking through the rookeries and counting the pups, a highly disruptive procedure. At smaller rookeries, with good vantage points, pups are occasionally counted from the periphery of rookeries without disturbing the sea lions. We evaluated counts made from medium-format, color, aerial photographs as an alternative to drive counts and peripheral counts. Neither the peripheral counts nor the aerial photographic counts disturbed animals on the rokeries. There were strong 1:1 linear relationships between photographic counts and drive counts ( r ²= 0.966, P < 0.001) and between photographic counts and peripheral counts ( r ²= 0.999, P < 0.001). Precision was similar for all three methods of counting. We suggest that medium-format, color, aerial photographs is appropriate for routine surveys of Steller sea lion pups in Alaska because it is not disruptive to the hauled-out sea lions and provides comparable estimates with similar precision to drive and peripheral counts. Large areas canbe rapidly surveyed during periods of good weather with a minimum of manpower.     

A dominant allele controls development into female mimic male and diminutive female ruffs

Maintaining polymorphisms for genes with effects of ecological significance may involve conflicting selection in males and females. We present data from a captive population of ruffs (Philomachus pugnax) showing that a dominant allele controls development into both small, ‘female mimic’ males (‘faeders’), and a previously undescribed class of small ‘female faeders’. Most male ruffs have elaborate breeding plumage and display behaviour, but 0.5–1.5% are faeders, which lack both. Females from a captive population previously lacking faeders were bred with two founder faeder males and their faeder sons. The faeders’ offspring had a quadrimodal size distribution comprising normal-sized males and females, faeders and atypically small females. By contrast, ornamented males fathered only normal-sized offspring. We conclude that both founding faeders were heterozygous for a faeder allele absent from the original population. This allele is dominant to previously described genes that determine development into independent versus satellite ornamented males. Unlike those genes, the faeder allele is clearly expressed in females. Small body size is a component of the male faeder mating strategy, but provides no obvious benefit to females. Bisexual expression of the gene provides the opportunity to quantify the strength of sexually antagonistic selection on a Mendelian trait.     

No Evidence of Metabolic Depression in Western Alaskan Juvenile Steller Sea Lions (Eumetopias jubatus)

Steller sea lion (Eumetopias jubatus) populations have undergone precipitous declines through their western Alaskan range over the last four decades with the leading hypothesis to explain this decline centering around changing prey quality, quantity, or availability for this species (i.e., nutritional stress hypothesis). Under chronic conditions of reduced food intake sea lions would conserve energy by limiting energy expenditures through lowering of metabolic rate known as metabolic depression. To examine the potential for nutritional stress, resting metabolic rate (RMR) and body composition were measured in free-ranging juvenile Steller sea lions (N = 91) at three distinct geographical locations (Southeast Alaska, Prince William Sound, Central Aleutian Islands) using open-flow respirometry and deuterium isotope dilution, respectively. Average sea lion RMR ranged from 6.7 to 36.2 MJ d⁻¹ and was influenced by body mass, total body lipid, and to a lesser extent, ambient air temperature and age. Sea lion pups captured in the Aleutian Islands (region of decline) had significantly greater body mass and total body lipid stores when compared to pups from Prince William Sound (region of decline) and Southeast Alaska (stable region). Along with evidence of robust body condition in Aleutian Island pups, no definitive differences were detected in RMR between sea lions sampled between eastern and western populations that could not be accounted for by higher percent total body lipid content, suggesting that that at the time of this study, Steller sea lions were not experiencing metabolic depression in the locations studied.     

The oldest known fur seal

The poorly known fossil record of fur seals and sea lions (Otariidae) does not reflect their current diversity and widespread abundance. This limited fossil record contrasts with the more complete fossil records of other pinnipeds such as walruses (Odobenidae). The oldest known otariids appear 5–6 Ma after the earliest odobenids, and the remarkably derived craniodental morphology of otariids offers few clues to their early evolutionary history and phylogenetic affinities among pinnipeds. We report a new otariid, Eotaria crypta, from the lower middle Miocene ‘Topanga’ Formation (15–17.1 Ma) of southern California, represented by a partial mandible with well-preserved dentition. Eotaria crypta is geochronologically intermediate between ‘enaliarctine’ stem pinnipedimorphs (16.6–27 Ma) and previously described otariid fossils (7.3–12.5 Ma), as well as morphologically intermediate by retaining an M₂ and a reduced M₁ metaconid cusp and lacking P_2–4 metaconid cusps. Eotaria crypta eliminates the otariid ghost lineage and confirms that otariids evolved from an ‘enaliarctine’-like ancestor.     

RISK OF EXTIRPATION OF STELLER SEA LIONS IN THE GULF OF ALASKA AND ALEUTIAN ISLANDS: A POPULATION VIABILITY ANALYSIS BASED ON ALTERNATIVE HYPOTHESES FOR WHY SEA LIONS DECLINED IN WESTERN ALASKA

We estimated the risk that the Steller sea lion will be extirpated in western Alaska using a population viability analysis (PVA) that combined simulations with statistically fitted models of historical population dynamics. Our analysis considered the roles that density‐dependent and density‐independent factors may have played in the past, and how they might influence future population dynamics. It also established functional relationships between population size, population growth rate and the risk of extinction under alternative hypotheses about population regulation and environmental variability. These functional relationships can be used to develop recovery criteria and guide research and management decisions. Life table parameters (e.g., birth and survival rates) operating during the population decline (1978–2002) were estimated by fitting simple age‐structured models to time‐series of pup and non‐pup counts from 33 rookeries (subpopulations). The PVA was carried out by projecting all 33 subpopulations into the future using these estimated site‐specific life tables (with associated uncertainties) and different assumptions about carrying capacities and the presence or absence of density‐dependent population regulation. Results suggest that the overall predicted risk of extirpation of Steller sea lions as a species in western Alaska was low in the next 100 yr under all scenarios explored. However, most subpopulations of Steller sea lions had high probabilities of going extinct within the next 100 yr if trends observed during the 1990s were to continue. Two clusters of contiguous subpopulations occurring in the Unimak Pass area in the western Gulf of Alaska/eastern Aleutian Islands and the Seguam–Adak region in the central Aleutian Islands had relatively lower risks of extinction. Risks of extinction for a number of subpopulations in the Gulf of Alaska were reduced if the increases observed since the late 1990s continue into the future. The risks of subpopulations going extinct were small when density‐dependent compensation in birth and survival rates was assumed, even when random stochasticity in these vital rates was introduced.     

Infectious disease and the decline of Steller sea lions (Eumetopias jubatus) in Alaska, USA: insights from serologic data

Serologic data were examined to determine whether infectious disease may have played a role in the decline of Steller sea lions (Eumetopias jubatus) in the Gulf of Alaska and Aleutian Islands, USA. Available published data, unpublished data, and recent collections (1997-2000) were compared and reviewed. Data were stratified by geography to compare the declining western Alaskan population in the Aleutian Islands through eastern Prince William Sound to the increasing population in southeastern Alaska. Prevalences of antibodies from the 1970s to the early 1990s were noted for Leptospira interrogans, Chlamydophila psittaci, Brucella spp., phocid herpesvirus-1, and calciviruses. Serum samples collected from 1997-2000 were tested for antibodies to these agents as well as to marine mammal morbilliviruses, canine parvovirus, and canine adenovirus-1 and -2. Conclusions could not be drawn about changes in antibody prevalence to these agents during the decline of Steller sea lions, however, because data were incomplete or not comparable as a result of inconsistencies in testing techniques. Despite these shortcomings, results provided no convincing evidence of significant exposure of Steller sea lions to morbilliviruses, Brucella spp., canine parvovirus, or L. interrogans. Steller sea lions have been exposed to phocid herpesviruses, caliciviruses, canine adenovirus, and C. psittaci or to cross-reactive organisms in regions of both increasing and decreasing sea lion abundance. Based on similar antibody prevalence estimates from the increasing and decreasing populations, these agents are unlikely to have been the primary cause of the population decline. They may have contributed to the decline or impeded population recovery, however, because of undetected mortality and morbidity or reductions of fecundity and body condition in animals under other stresses. Systematic monitoring for disease agents and their effects is needed to determine whether infectious disease currently plays a role in the decline and lack of recovery of Steller sea lions.     

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.     

Effects of research disturbance on the behavior and abundance of Steller sea lions (Eumetopias jubatus) at two rookeries in Alaska

We examined the effects of research disturbance on the behavior and abundance of Steller sea lions (Eumetopias jubatus) at rookeries on Marmot and Ugamak Islands in Alaska. During 3 of 6 yr, researchers intentionally drove all adult and juvenile sea lions off at least part of the beach in order to permanently mark and measure sea lion pups. The research disturbance occurred after the majority of females had bred and when most pups were 1 mo old. We used generalized linear models to determine the relationship between research disturbance and sea lion behavior or abundance. Research disturbance was related to changes in the proportion of sea lions exhibiting two to three of nine behavior metrics: agonistic and resting females and active males at Marmot, and active and resting males and females at Ugamak. Model results indicated that changes lasted between 3 and 20 d depending on the sex, behavior, and rookery. Inclusion of research disturbance into Marmot abundance models did not improve the fit to the data, if variability between years was permitted. Optimally timed, low‐frequency research disturbance did not appear to have long‐term effects on sea lion behavior or abundance and was largely associated with changes that were similar to natural variation.     

Finding the ‘lost years’ in green turtles: insights from ocean circulation models and genetic analysis

Organismal movement is an essential component of ecological processes and connectivity among ecosystems. However, estimating connectivity and identifying corridors of movement are challenging in oceanic organisms such as young turtles that disperse into the open sea and remain largely unobserved during a period known as ‘the lost years’. Using predictions of transport within an ocean circulation model and data from published genetic analysis, we present to our knowledge, the first basin-scale hypothesis of distribution and connectivity among major rookeries and foraging grounds (FGs) of green turtles (Chelonia mydas) during their ‘lost years’. Simulations indicate that transatlantic dispersal is likely to be common and that recurrent connectivity between the southwestern Indian Ocean and the South Atlantic is possible. The predicted distribution of pelagic juvenile turtles suggests that many ‘lost years hotspots’ are presently unstudied and located outside protected areas. These models, therefore, provide new information on possible dispersal pathways that link nesting beaches with FGs. These pathways may be of exceptional conservation concern owing to their importance for sea turtles during a critical developmental period.     

Trophic level and overlap of sea lions (Zalophus californianus) in the Gulf of California, Mexico

Stable isotope and scat analyses were used in concert to determine trophic level and dietary overlap among California sea lions from different rookeries in the Gulf of California. Isotopic analysis of the fur of sea lion pups revealed differences in δ¹⁵N and δ¹³C values among rookeries during the breeding season. Mean δ¹⁵N and δ¹³C values varied from 20.2‰ to 22.4‰ and from −15.4‰ to −14.0‰, respectively. The pattern of differences among rookeries was similar between years in most cases. Isotopic variations among rookeries were associated with differences in prey consumption. There was a significant correlation between δ¹⁵N value and trophic level, as determined by scat analysis. Joint application of isotopic and scat analyses allowed us to identify how the feeding habits of sea lions vary with location. Our results suggest the presence of spatial structure in available prey as well as the localized use of prey by sea lions across the Gulf of California.     

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.