Mark‐recapture modeling accounting for state uncertainty provides concurrent estimates of survival and fecundity in a protected harbor seal population

Harbor seal breeding behavior and habitats constrain opportunities for individual‐based studies, and no current estimates of both survival and fecundity exist for any of the populations studied worldwide. As a result, the drivers underlying the variable trends in abundance exhibited by harbor seal populations around the world remain uncertain. We developed an individual‐based study of harbor seals in northeast Scotland, whereby data were collected during daily photo‐identification surveys throughout the pupping seasons between 2006 and 2011. However, a consequence of observing seals remotely meant that information on sex, maturity‐stage, or breeding status was not always available. To provide unbiased estimates of survival rates we conditioned initial release of individuals on the first time sex was known to estimate sex‐specific survival rates, while a robust design multistate model accounting for uncertainty in breeding status was used to estimate reproductive rate of multiparous and ≥3‐yr‐old females. Survival rates were estimated at 0.95 (95% CI = 0.91–0.97) for females and 0.92 (0.83–0.96) for males, while reproductive rate was estimated at 0.89 (0.75–0.95) for multiparous and 0.69 (0.64–0.74) for ≥3‐yr‐old females. Stage‐based population modeling indicated that this population should be recovering, even under the current shooting quotas implemented by the recent management plan.     

Brain size in neonatal and adult Weddell seals: Costs and consequences of having a large brain

Little is known about the ontogeny of brain size in pinnipeds despite potential functional implications of brain substrate (glucose, oxygen) requirements for diving, fasting, growth, and lactation strategies. We measured brain mass (brM) and cranial capacity (CC) in newborn and adult Weddell seals. Neonatal Weddell seals had brM that represented ~70% of adult brM. Weddell seals have the largest neonatal brain, proportional to adult brain, reported for any mammal to date, which is remarkable considering the relatively small size of Weddell seal pups at birth (6%–7% of maternal body mass) compared to neonates of other highly precocial mammals. Provision of sufficient glucose to maintain the large, well‐developed brain of the neonatal Weddell seal has a nontrivial metabolic cost to both pup and mother. We therefore hypothesize that this phenomenon must have functional significance, such as allowing pups to acquire complex under‐ice navigation skills during the period of maternal attendance.     

Trend changes in sympatric Subantarctic and Antarctic fur seal pup populations at Marion Island,Southern Ocean

Recent pup population estimates of sympatric Subantarctic (Arctocephalus tropicalis) and Antarctic fur seals (A. gazella) at Marion Island are presented. Published pup population estimates of A. tropicalis (1995 and 2004) with an unpublished total island count in 2013, and annual counts on subsets of rookeries (2007–2015) were analyzed using a hierarchical Bayesian model. The pup population declined by 46% (95% credible interval CI: 43%–48%) between 2004 (mean = 15,260, CI: 14,447–16,169 pups) and 2013 (mean = 8,312, CI: 7,983–8,697), mirrored by a 58%–60% decline at rookeries counted annually (2007–2015). Population decline was highest at high‐density west and north coast rookeries, despite negligible change in female attendance patterns, pup mortality or median pupping date over the previous 25 yr. A better understanding of foraging behavior and its effects on reproductive success and survival in this A. tropicalis population is needed before we can attribute population decline to any external factors. In contrast, total island counts of A. gazella pups in 2007, 2010, and 2013, suggest that this population is still increasing although the annual intrinsic rate of population growth decreased from 17.0% (1995–2004, 744 pups) to 4.0% (2010–2013, 1,553 pups). The slowed growth of A. gazella is likely the result of saturation at the main rookery.     

Variation in breeding phenology provides insights into drivers of long-term population change in harbour seals

Phenological trends provide important indicators of environmental change and population dynamics. However, the use of untested population-level measures can lead to incorrect conclusions about phenological trends, particularly when changes in population structure or density are ignored. We used individual-based estimates of birth date and lactation duration of harbour seals (Phoca vitulina) to investigate energetic consequences of changes in pupping phenology. Using generalized linear mixed models, we first demonstrate annual variation in pupping phenology. Second, we show a negative relationship between lactation duration and the timing of pupping, indicating that females who pup early nurse their pups longer, thereby highlighting lactation duration as a useful proxy of female condition and resource availability. Third, individual-based data were used to derive a population-level proxy that demonstrated an advance in pupping date over the last 25 years, co-incident with a reduction in population abundance that resulted from fisheries-related shootings. These findings demonstrate that phenological studies examining the impacts of climate change on mammal populations must carefully control for changes in population density and highlight how joint investigations of phenological and demographic change provide insights into the drivers of population declines.     

Primiparous females do not exhibit reduced maternal care in gray seals (Halichoerus grypus)

We compared the behaviors of primiparous and multiparous gray seal (Halichoerus grypus) females over the course of lactation to examine whether poorly developed maternal behaviors may play a role in the reduced lactation performance observed in primiparous females. Overall, primiparous females spent as much time interacting with their pups as multiparous females. The proportion of time spent nursing their pup increased significantly between early and peak lactation in both primiparous and multiparous females. Although there was no significant difference in the duration of nursing bouts as a function of reproductive status, primiparous females nursed significantly more frequently (bouts/hour) and, therefore, spent a significantly greater proportion of time nursing than multiparous females throughout lactation. Primiparous gray seal females were also significantly more active than multiparous females, however, the difference in activity represented only a small proportion of the overall time budget. We conclude that poorly developed maternal behaviors resulting from a lack of prior reproductive experience are unlikely to account for lower levels of milk energy transfer to pups in primiparous gray seals.     

A longitudinal and cross-sectional analysis of total body oxygen store development in nursing harbor seals (<Emphasis Type="Italic">Phoca vitulina</Emphasis>)

This study compared the efficacy of longitudinal and cross-sectional sampling regimes for detecting developmental changes in total body oxygen (TBO₂) stores that accompany behavioral development in free-ranging harbor seal pups. TBO₂ stores were estimated for pup (n = 146) and adult female (n = 20) harbor seals. Age related changes were compared between pups captured repeatedly during the lactation period (longitudinal dataset) and a second group of pups handled only once (cross-sectional dataset). At each handling, hematocrit, hemoglobin, red blood cell count, total plasma volume, blood volume, muscle myoglobin concentration, and blood and muscle oxygen stores were determined. Comparisons across age categories revealed newborn blood oxygen stores were initially elevated, declined to low values by early lactation, and increased through post-weaning. Muscle oxygen stores remained low and constant throughout lactation and only increased significantly post-weaning. Overall TBO₂ stores increased 17% during lactation, and weaned pups had TBO₂ stores that were 55% as large as those of adults. Thus, significant increases in TBO₂ stores must occur after weaning, as pups begin to forage independently. Results from the two sampling schemes did not differ, indicating that the logistically simpler cross-sectional design can be used to monitor physiological development in harbor seals.     

Influence of maternal mass and condition on energy transfer in Weddell seals

1. Environmental variation influences food abundance and availability, which is reflected in the reproductive success of top predators. We examined maternal expenditure, offspring mass and condition for Weddell seals in 2 years when individuals exhibited marked differences in these traits. 2. For females weighing > or = 355 kg there was a positive relationship between maternal post-partum mass (MPPM) and lactation length, but below this there was no relationship, suggesting that heavier females were able to increase lactation length but lighter females were restricted to a minimum lactation period of 33 days. 3. Overall, females were heavier in 2002, but in 2003 shorter females were lighter than similar-sized females in 2002 suggesting that the effects of environmental variability on foraging success and condition are more pronounced in smaller individuals. 4. There was no relationship between MPPM and pup birth mass, indicating pre-partum investment did not differ between years. However, there was a positive relationship between MPPM and pup mass gain. Mass and energy transfer efficiency were 10.2 and 5.4% higher in 2002 than 2003, which suggests costs associated with a putatively poor-resource year were delayed until lactation. 5. Heavier females lost a higher proportion of mass during lactation in both years, so smaller females may not have been able to provide more to their offspring to wean a pup of similar size to larger females. 6. MPPM had only a small influence on total body lipid; therefore, regardless of mass, females had the same relative body composition. Females with male pups lost a higher percentage of lipid than those with female pups, but by the end of lactation female pups had 4.5% higher lipid content than males. 7. It appears that for Weddell seals the consequences of environmentally induced variation in food availability are manifested in differences in maternal mass and expenditure during lactation. These differences translate to changes in pup mass and condition at weaning with potential consequences for future survival and recruitment.     

Variation in milk production and lactation performance in grey seals and consequences for pup growth and weaning characteristics

Phocid seals are one of the few groups of mammals capable of sustaining the energetic demands of lactation entirely through body nutrient stores while fasting. Lactation performance of the female in turn influences the rate and pattern of pup growth. We examined variation in and patterns of milk composition and production, maternal energy output, and pup growth and energy deposition over the entire lactation period in 18 grey seal mother-pup pairs using hydrogen isotope (3H2O and D2O) dilution. Milk composition was independent of maternal mass and nutrient stores, indicating dependence on other physiological and genetic factors. Heavier females lactated longer (r2=0.653, P<0.001), had higher total milk outputs (r2=0.652, P<0.001), and produced larger pups at weaning (r2=0.417, P=0.005). While fatter females lactated for longer periods of time (r2=0.595, P<0.001), females with a larger lean body mass at parturition produced more milk (r2=0.579, P<0.001). Total milk energy output was the strongest predictor of pup weaning mass, which, along with the pup's efficiency of energy storage, accounted for 91% of the variation in weaning mass. Nevertheless, there was sufficient plasticity in milk composition and energy output that some smaller females produced relatively large pups. Few females appeared to deplete body nutrients to the point where it might limit the duration of lactation.     

The influence of reproductive experience on milk energy output and lactation performance in the grey seal (Halichoerus grypus)

Although evidence from domestic and laboratory species suggests that reproductive experience plays a critical role in the development of aspects of lactation performance, whether reproductive experience may have a significant influence on milk energy transfer to neonates in wild populations has not been directly investigated. We compared maternal energy expenditures and pup growth and energy deposition over the course of lactation between primiparous and fully-grown, multiparous grey seal (Halichoerus grypus) females to test whether reproductive experience has a significant influence on lactation performance. Although there was no difference between primiparous females in milk composition and, thus, milk energy content at either early or peak lactation primiparous females had a significantly lower daily milk energy output than multiparous females indicating a reduced physiological capacity for milk secretion. Primiparous females appeared to effectively compensate for lower rates of milk production through an increased nursing effort and, thus, achieved the same relative rate of milk energy transfer to pups as multiparous females. There was no difference between primiparous and multiparous females in the proportion of initial body energy stores mobilised to support the costs of lactation. Although primiparous females allocated a greater proportion of energy stores to maternal maintenance versus milk production than multiparous females, the difference was not sufficient to result in significant differences in the efficiency of energy transfer to pups. Thus, despite a lower physiological capacity for milk production, primiparous females weaned pups of the same relative size and condition as multiparous females without expending proportionally more energy. Although reproductive experience does not significantly affect the overall lactation performance of grey seals, our results suggest that increases in mammary gland capacity with reproductive experience may play a significant role in the age-related increases in neonatal growth rates and weaning masses observed in other free-ranging mammals.     

Postrelease movement of rehabilitated harbor seal (Phoca vitulina richardii) pups compared with cohort‐matched wild seal pups

Harbor seal (Phoca vitulina richardii) populations in the inland waters of Washington and British Columbia are at or near carrying capacity. Stranded pups often are collected and admitted to rehabilitation centers, and then released when they reach a weight of 22 kg and meet a variety of preestablished health and release conditions. While rehabilitation is common practice, it is unclear if rehabilitated seal pups behave like wild weaned pups. Using satellite transmitters, we compared movement patterns of 10 rehabilitated pups with 10 wild weaned pups. When released, rehabilitated seals were longer and heavier than wild pups, while wild pups had a larger mean axillary girth. No clinically different blood parameters were detected. On average, rehabilitated harbor seal pups traveled nearly twice as far cumulatively, almost three times as far daily, and dispersed over three times as far from the release site compared to wild weaned seals. Additionally, wild harbor seals transmitted nearly twice as long as did rehabilitated seals. These patterns suggest that learned behavior during the brief 3–4 wk nursing period likely enables wild harbor seal pups to move less daily and remain closer to their weaning site than rehabilitated pups.     

Grey seal, Halichoerus grypus, energetics: females invest more in male offspring

Grey seal females transfer large amounts of energy to their pups during the brief lactation period. The costs of lactation have been measured using weight changes of mother and pup pairs. Large females come ashore to give birth earlier in the season and lose weight more rapidly than smaller females. The sex ratio of Grey seal pups born is skewed towards males in the early part of the breeding season. Male pups are larger at birth and gain weight more rapidly than female pups, and their mothers show a correspondingly faster rate of weight loss than mothers of female pups. The energy costs of gestation and lactation to a Grey seal mother are 31 GJ for male pups and2–8 GJ for female pups. Males are therefore 10% more costly in energy terms to raise to weaning. Because, on average, large females arrive at breeding sites before smaller animals, biased results on weight changes would be obtained from methods which do not use repeated weighings. We suggest that the high efficiencies of lactation estimated for Harp seals compared with other phocid seals could be accounted for by such a bias.     

Reaction of Harbor Seals to Cruise Ships

Abstract: The largest aggregations of harbor seals (Phoca vitulina) in Alaska, USA, haul out on floating ice in tidewater glacial fjords. Seals use these fjords in peak numbers during the critical periods of pupping, breeding, and molting when visits by tour ships also peak. Documented and suspected declines of harbor seals in fjords with rising vessel traffic underscore the need to better understand possible impacts, particularly in areas where ship visits have risen substantially in the past 2 decades. We examined the interruption of haul-out bouts of harbor seals due to approaching cruise ships in Disenchantment Bay, Alaska. We conducted observations from cruise ships and focused on disturbance of seals as evidenced by seals flushing into the water from the floating ice on which they rested. We investigated rate of flushing in relation to vessel distance, approach angle, group size, and seal type (mother, pup, or other). Using a survival-regression analysis, we found that the risk of disturbing harbor seals increased when ships approached within 500 m; seals approached as close as 100 m were 25 times more likely to enter the water than seals 500 m from a ship. Seals were 4 times more prone to enter the water when ships were approaching directly rather than passing abeam. Seals responded similarly regardless of group size or seal type. Energetic models indicated a potential to disrupt energy balance and cause thermal stress in disturbed pups if they spent >50% of their time in ice-chilled water. Studies at non-glacial sites suggest that pups spend 40–70% of their time in the water. Voluntary guidelines for approaching seals in Alaska recommend that cruise ships approach ≥91 m (100 yards), a distance at which we show 90% of seals would flush into the water. Our findings indicate a need to develop regulations to maintain a 500-m separation between cruise ships and seals in all Alaskan glacial fjords.     

Feast or famine: evidence for mixed capital–income breeding strategies in Weddell seals

Evolved patterns of resource expenditure for reproduction have resulted in a life history continuum across species. A strictly capital-breeding strategy relies extensively on stored energy for reproduction, whereas income breeding uses energy acquired throughout the reproductive period. However, facultative income breeding has been shown in some classically capital-breeding animals, and was originally thought to provide a nutritional refuge for smaller females incapable of securing sufficient reserves during pre-partum foraging. We examined milk composition and milk output for the Weddell seal to determine to what degree lactation was aided by food intake, and what factors contributed to its manifestation. Milk composition was independent of maternal post-partum mass and condition, but did change over lactation. Changes were most likely in response to energetic and nutritional demands of the pup at different stages of development. During early lactation, females fasted and devoted 54.9% of total energy loss to milk production. Later in lactation 30.5% more energy was devoted to milk production and evidence suggested that larger females fed more during lactation than smaller females. It appears that Weddell seals may exhibit a flexible strategy to adjust reproductive investment to local resource levels by taking advantage of periods when prey are occasionally abundant, although it is restricted to larger females possessing the physiological capacity to dive for longer and exploit different resources during lactation. This supports the assumption that although body mass and phylogenetic history explain most of the variation in lactation patterns (20–69%), the remaining variation has likely resulted from physiological adaptations to local environmental conditions. Our study confirms that Weddell seals use a mixed capital–income breeding strategy, and that considerable intraspecific variation exists. Questions remain as to the amount of energy gain derived from the income strategy, and the consequences for pup condition and survival. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Lactating Females Do Not Discriminate Between Their Own Young and Unrelated Pups in the Communally Breeding Rodent, Octodon degus

Females in numerous rodent species engage in communal nesting and breeding, in which they share one or more nests to rear their young. A potential cost of communal nesting and breeding is that mothers divert resources to unrelated offspring. One way mothers could avoid this cost is to recognize and favour their own young over unrelated offspring when allocating maternal effort. We assessed whether female degus (Octodon degus), a communally nesting and breeding caviomorph rodent, discriminate between their own and unrelated offspring during lactation. Female degus previously have been shown to distinguish between their own and unrelated pups when exposed to odours from both. We measured pup discrimination based on differences in the retrieval behaviour of females that were in early or intermediate lactation directed towards their own and unrelated offspring; offspring presented were of similar or different age. Before any event of pup retrieval, lactating females spent similar amounts of time and interacted to a similar extent with their own and unrelated pups. During pup retrieval, all lactating females transported both pups to the nest. Neither relatedness to pups, nor pup‐age differences, influenced the order in which pups were retrieved to the nest. Dams waited similar amounts of time before retrieving the first pup when the first transported young was their own or unrelated. Likewise, females waited similar amounts of time before retrieving the second pup when the pup transported first was their own or unrelated. The time between first and second pup transport events was longer when dams were in early when compared with intermediate lactation, but only when pups were of similar age. All experimental subjects nursed unrelated pups after they were retrieved. Collectively, our results do not support the hypothesis that communally breeding female degus use their recognition ability to discriminate against unrelated offspring in favour of their own young.     

Age-related reproductive variation in a wild marine mammal population

Life history theory predicts a change in reproduction success with age as energy resources are limited and must be allocated effectively to maximize reproduction and survival. In this study, we use three reproductive performance measures, maternal expenditure, offspring weaning mass, and first-year survival, to investigate the role that maternal age plays in successful reproduction. Long-term uninterrupted life history data available for Marion Island’s southern elephant seals and mass change estimates from photogrammetry data allow for assessment of age-related reproduction performance and trade-offs. Known-aged adult females were photographed for photogrammetric mass estimation (n = 29) and their pups weighed at weaning during the 2009 breeding season. Maternal age and proportional mass loss positively influenced pup weaning mass. In turn, first-year pup return rates (as a proxy for survival) were assessed through the intensive mark–recapture program. Pup survival increased with female age and weaning mass. Pups of young females aged 3–6 years have a lower first-year survival probability compared with pups of older and larger females.     

REPRODUCTIVE PATTERNS OF THE HAWAIIAN MONK SEAL

We evaluated reproductive patterns of the Hawaiian monk seal ( Monachus schauinslandi ) using a combination of fitted age-specific reproductive curves and analysis of reproductive patterns of individual females. We review the difficulties inherent in the acquisition and modeling of reproductive data with emphasis on the significance of reproductive senescence to populations with dissimilar age/sex compositions. Validation of the fitted reproductive parameters was accomplished by Monte Carlo sampling of parameter distributions to compare the expected number of pups with the observed production. Although the fitted reproductive functions appear to provide an acceptable fit to the raw reproductive data, we found that the fitted curves did a poor job of predicting the actual pup production in individual years because of high variability among years. To further verify, and elaborate on, the patterns in the pooled (multi-seal, and multi-year) rates, we examined attributes of the reproductive performance of individual seals. The attributes included age of primiparity, reproductive rates computed over several age ranges, and the relationship between reproductive performance and seal longevity. Analysis of individual seal patterns reinforced the conclusion that reproductive senescence is operative in monk seal populations.     

Sighting patterns reveal unobserved pupping events to revise reproductive rate estimates for Hawaiian monk seals in the main Hawaiian Islands

We used sighting reports, including decades of citizen-reported Hawaiian monk seal (Neomonachus schauinslandi) sightings, to describe female breeding biology and reproductive success in the main Hawaiian Islands. We first used this data set to describe the timing of events in the female reproductive cycle. We then conducted an expert review of patterns in sighting histories to detect unobserved pupping events. Finally, we estimated the age-specific reproductive curve for female monk seals in the main Hawaiian Islands. Charting reproductive cycles showed indications of the robust condition of female monk seals in the main Hawaiian Islands; they nursed pups 12% longer than their counterparts in the Northwestern Hawaiian Islands and regained condition to molt more quickly after weaning a pup. By examining sighting histories, we were able to infer 25 unobserved pupping events that had previously gone uncounted. We accounted for additional uncertainty with a randomization procedure. After accounting for unobserved pupping events, the age-specific reproductive rate of main Hawaiian Islands monk seals exceeded 0.70 for prime aged females (8–18 years). This is the highest reproductive rate reported for any of the Hawaiian monk seal breeding sites, illustrating the important role of the main Hawaiian Islands population in Hawaiian monk seal recovery.     

Postbranding Survival of Steller Sea Lion Pups at Lowrie Island in Southeast Alaska

ABSTRACT Steller sea lion (Eumetopias jubatus) pups (n = 366) were hot-branded at Lowrie Island, Southeast Alaska, USA, in June 2001 and 2002 for vital-rates studies. To assess potential mortality following branding, we estimated weekly survival to 12 weeks postbranding using mark-recapture models. Survival estimates ranged from 0.984/week to 0.988/week, or 0.868 over the 12-week period; varied little with sex, year, and capture area; and were higher for larger than smaller male pups and unexpectedly lower for larger than smaller female pups. Inclusion of resights at 1–3 years of age prevented a −4.5% bias in cumulative survival to 12 weeks postbranding by accounting for pups that survived but permanently emigrated from Lowrie Island during the 12-week survey. Data from double-marked pups (i.e., branded and flipper-tagged) indicated the low brand-misreading probability of 3.1% did not bias survival estimates. Assuming survival differences between the first 2 weeks postbranding and later weeks were due entirely to the branding event, potential postbranding mortality of branded pups attributable to the branding event was 0.5–0.7%, or one pup for every 200 marked. Weekly survival of branded pups was nearly identical to estimates from a control group of undisturbed, unbranded pups born to 10–11-year-old branded adult females in 2005 (0.987–0.988/week) and similar to pup survival estimates from other otariid studies. Available data did not indicate substantial mortality to 12 weeks postbranding resulting from the branding disturbance, suggesting branding of Steller sea lion pups can be used effectively for investigations of population declines without significantly affecting population health or study goals.     

ESTIMATING PUP PRODUCTION OF HARP SEALS, PAGOPHILUS GROENLANDICUS, IN THE NORTHWEST ATLANTIC

Photographic and visual aerial surveys to determine current pup production of Northwest Atlantic harp seals were conducted off Newfoundland and in the Gulf of St. Lawrence during March 1999-Photographic surveys were conducted on all whelping concentrations between 14 and 24 March, whereas a visual survey was made of the southern Gulf concentrations on 14 March. Pup production was estimated to be 739,100 (SE = 96,300, CV = 13.0%) at the Front, 82,600 (SE = 22,500, CV = 27.2%) in the northern Gulf, and 176,200 (SE = 25,400, CV = 14.4%) in the southern Gulf (Magdalen Island) for a total of 997,900 (SE = 102,100, 10.2%). Changes in aerial survey estimates indicate that pup production has increased since 1994. A new method to correct for the temporal change in the proportion of pups present on the ice was examined by fitting the percentage of pups observed in three age-dependent stages to a Normal distribution. The results were compared to those obtained from a more complex model used previously. The Simple model produced slightly higher, and hence more conservative, estimates of the proportion of births that had occurred before the time of the survey than the Complex model. When using the Simple model fewer assumptions regarding the start date of pupping and the proportion of older pups remaining on the ice were required, the herd had to be followed for a shorter period, and a more convenient means of calculating confidence limits was available.     

DECLINES IN HARBOR SEAL (PHOCA VITULINA) NUMBERS IN GLACIER BAY NATIONAL PARK, ALASKA, 1992–2002

Glacier Bay National Park had one of the largest breeding aggregations of harbor seals in Alaska, and it is functionally the only marine reserve for harbor seals in Alaska; yet, numbers of seals in the Bay are declining rapidly. Understanding why seals in Glacier Bay are declining may clarify their minimal habitat needs. We estimated population trends using models that controlled for environmental and observer‐related factors. In 1992, 6,200 seals were counted on icebergs in a tidewater glacial fjord and at terrestrial sites; by 2002 only 2,550 seals were counted at these same haul‐outs. Numbers of non‐pups in the glacial fjord declined by 6.6%/yr (?39%/8 yr) in June and by 9.6%/yr (?63%/11 yr) in August and at all other haul‐outs by 14.5%/yr (?75%/10 yr) during August. In the glacial fjord the number of pups remained steady from 1994 to 1999 and made up an increasing proportion of seals counted (5.4%/yr), and the proportion of pups peaked at 34%–36%. The rapid declines do not appear to be due to changes in seal behavior or redistribution. The declines reinforce genetic evidence that harbor seals in Glacier Bay are demographically isolated from other populations and indicate that current management stocks need to be redefined. Changes in Glacier Bay's ecosystem and population demographic data from the glacial fjord suggest that interspecific competition and predation are likely factors in the declines.