Age-specific reproduction in female Steller sea lions in Southeast Alaska.

Age-, region-, and year-specific estimates of reproduction are needed for monitoring wildlife populations during periods of ecosystem change. Population dynamics of Steller sea lions (Eumetopias jubatus) in Southeast Alaska varied regionally (with high population growth and survival in the north vs. the south) and annually (with reduced adult female survival observed following a severe marine heatwave event), but reproductive performance is currently unknown. We used mark-resighting data from 1006 Steller sea lion females marked as pups at ~3 weeks of age from 1994 to 1995 and from 2001 to 2005 and resighted from 2002 to 2019 (to a maximum age of 25) to examine age-, region-, and year-specific reproduction. In the north versus the south, age of first reproduction was earlier (beginning at age 4 vs. age 5, respectively) but annual birth probabilities of parous females were reduced by 0.05. In an average year pre-heatwave, the proportion of females with pup at the end of the pupping season peaked at ages 12-13 with ~0.60/0.65 (north/south) with pup, ~0.30/0.25 with juvenile, and ~0.10 (both regions) without a dependent. In both regions, reproductive senescence was gradual after age 12: ~0.40, 0.40, and 0.20 of females were in these reproductive states, respectively, by age 20. Correcting for neonatal mortality, true birth probabilities at peak ages were 0.66/0.72 (north/south). No cost of reproduction on female survival was detected, but pup production remained lower (-0.06) after the heatwave event, which if sustained could result in population decline in the south. Reduced pup production and greater retention of juveniles during periods of poor prey conditions may be an important strategy for Steller sea lions in Southeast Alaska, where fine-tuning reproduction based on nutritional status may improve the lifetime probability of producing pups under good conditions in a variable and less productive environment.

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.

Development and application of DNA techniques for validating and improving pinniped diet estimates.

Polymerase chain reaction techniques were developed and applied to identify DNA from >40 species of prey contained in fecal (scat) soft-part matrix collected at terrestrial sites used by Steller sea lions (Eumetopias jubatus) in British Columbia and the eastern Aleutian Islands, Alaska. Sixty percent more fish and cephalopod prey were identified by morphological analyses of hard parts compared with DNA analysis of soft parts (hard parts identified higher relative proportions of Ammodytes sp., Cottidae, and certain Gadidae). DNA identified 213 prey occurrences, of which 75 (35%) were undetected by hard parts (mainly Salmonidae, Pleuronectidae, Elasmobranchii, and Cephalopoda), and thereby increased species occurrences by 22% overall and species richness in 44% of cases (when comparing 110 scats that amplified prey DNA). Prey composition was identical within only 20% of scats. Overall, diet composition derived from both identification techniques combined did not differ significantly from hard-part identification alone, suggesting that past scat-based diet studies have not missed major dietary components. However, significant differences in relative diet contributions across scats (as identified using the two techniques separately) reflect passage rate differences between hard and soft digesta material and highlight certain hypothesized limitations in conventional morphological-based methods (e.g., differences in resistance to digestion, hard part regurgitation, partial and secondary prey consumption), as well as potential technical issues (e.g., resolution of primer efficiency and sensitivity and scat subsampling protocols). DNA analysis of salmon occurrence (from scat soft-part matrix and 238 archived salmon hard parts) provided species-level taxonomic resolution that could not be obtained by morphological identification and showed that Steller sea lions were primarily consuming pink (Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon. Notably, DNA from Atlantic salmon (Salmo salar) that likely originated from a distant fish farm was also detected in two scats from one site in the eastern Aleutian Islands. Overall, molecular techniques are valuable for identifying prey in the fecal remains of marine predators. Combining DNA and hard-part identification will effectively alleviate certain predicted biases and will ultimately enhance measures of diet richness, fisheries interactions (especially salmon-related ones), and the ecological role of pinnipeds and other marine predators, to the benefit of marine wildlife conservationists and fisheries managers.

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.

Serum chemistry reference ranges for Steller sea lion (Eumetopias jubatus) pups from Alaska: stock differentiation and comparisons within a North Pacific sentinel species.

Blood chemistry and hematologic reference ranges are useful for population health assessment and establishing a baseline for future comparisons in the event of ecosystem changes due to natural or anthropogenic factors. The objectives of this study were to determine if there was any population spatial structure for blood variables of Steller sea lion (Eumetopias jubatus), an established sentinel species, and to report reference ranges for appropriate populations using standardized analyses. In addition to comparing reference ranges between populations with contrasting abundance trends, data were examined for evidence of disease or nutritional stress. From 1998 to 2011, blood samples were collected from 1,231 pups captured on 37 rookeries across their Alaskan range. Reference ranges are reported separately for the western and eastern distinct population segments (DPS) of Steller sea lion after cluster analysis and discriminant function analysis (DFA) supported underlying stock structure. Variables with greater loading scores for the DFA (creatinine, total protein, calcium, albumin, cholesterol, and alkaline phosphatase) also were greater for sea lions from the endangered western DPS, supporting previous studies that indicated pup condition in the west was not compromised during the first month postpartum. Differences between population segments were likely a result of ecological, physiological, or age related differences.