BREEDING BIOLOGY OF SOUTHERN ELEPHANT SEALS IN PATAGONIA

Abstract: Elephant seals breed in Patagonia (Península Valdés, Argentina) from late August to early November, reaching peak numbers during the first week in October. Observations of this population over the past ten years yielded similar results. Eighty percent of the pups were born by 2 October. Most (96%) of 663 females marked during three breeding seasons gave birth to a pup. Females stayed on land a mean of 28 d, gave birth 6 d after arrival, nursed their pups for 22 d, and copulated a mean of 2.5 times 20 d after parturition and 2 d before departure. Copulations peaked during the third week in October. Males spent 57–80 d on land fasting and defending harems of up to 134 females (median 11–13 females, depending on year). Most (96%) marked females that gave birth ( n = 636) also weaned their pups successfully. Pup sex ratio was unity. Harems were smaller and breeding occurred about three weeks earlier in Patagonia than in other colonies. Thermal conditions, day length and food availability may explain clines in the timing of breeding events between populations, Other parameters of the breeding season for the expanding Patagonia colony are similar to those for declining southern elephant seal populations elsewhere.     

Birth timing and population changes in greater horseshoe bat colonies (Rhinolophus ferrumequinum) are synchronized by climatic temperature

We counted the births of greater horseshoe bats born at three small breeding colonies in southwest Wales and south-west England at the extreme edge of the species distribution over nine summers (1984–1993). Overall birth timing was almost identical at the three sites (mean of mean birth dates = 13–14 July), but varied widely from year to year. Mean birth timing was synchronized at the three sites in a given year. Early births followed warm springs; a significant negative regression of mean birth date on mean April plus May temperature was evident. A rise of 2^oC accelerated mean birth date by about 18 days. Population levels fell at all three sites following the very late mean birth date of 28 July recorded in 1986, after an extremely cold spring and summer. Recovery of populations in all sites followed a series of warm springs and early mean birth dates, but was hampered by birth sex ratios favouring males for several years. These findings confirm climate, through its effect upon birth timing, and possibly also on sex ratios, as a major factor controlling bat populations, and hence the distribution of bat species.     

BODY TEMPERATURE IN THE NEWBORN SOUTHERN ELEPHANT SEAL, MIROUNGA LEONINA, AT MACQUARIE ISLAND

Newborn southern elephant seal pups were reported by Laws (1953) to be "to some extent poikilothermic at birth." Rectal temperatures of known age southern elephant seal pups were recorded during the 1985 pupping season at Macquarie Island. The mean pup rectal temperature was found to be 381°C ± 0.1°C SEM ( n = 131, range = 36.5°-39.1°C). Pups at two hours, six hours, and one day after birth had significantly higher rectal temperatures than pups two, three, or four days of age. Rectal temperatures of neonatal southern elephant seals were within the range observed for other pinnipeds, (but never as low as the 31°C previously observed for southern elephant seals at Signy Island in 1953). A significant though weak positive correlation was found between pup temperature and body weight. However, no correlation between pup temperature and age or any environmental factor was found. These observations demonstrated that southern elephant seal pups at Macquarie Island are homeothermic, rather than heterothermic from birth.     

THE INFLUENCE OF CLIMATIC SEASONALITY ON THE LIFE CYCLE OF THE PRIBILOF NORTHERN FUR SEAL

Weather conditions recorded from 1956 to 1986 on St. Paul Island, Alaska, were probed to establish their influence upon the northern fur seal's life cycle ( Callorhinus ursinus ). Air temperatures, wind speeds, and relative humidity levels were seasonally decomposed and compared with the timing of pupping and migration. Most pups are born in early July when air temperatures and relative humidity approach their highest annual levels and wind speeds are at their lowest. Weather conditions favor growth and survival of pups from July to September but are unfavorable in June. A rapid deterioration in weather through October and November corresponds with the fall migration of pups and lactating females. The data suggest the pivotal event in the fur seal's life cycle is the timing of birth and survival of nursing pups. As such, the ultimate determinant of the precisely timed fur seal life cycle appears to be climatic seasonality during the breeding season.     

Photo-ID-based estimates of reproductive patterns in female harbor seals

Variations in reproductive patterns offer important insights into the dynamics of pinniped populations, but collecting data on reproduction for species that spend much of the breeding season in the water is problematic. We used land‐based photo‐identification techniques to collect individual‐based data on the timing of pupping, total pup production, and lactation duration in a population of harbor seals in NE Scotland. Capture–Mark–Recapture (CMR) techniques were used to overcome potential biases due to changes in probability of capture, and provide estimates of lactation duration based upon changes in the “survival” of mother–pup bonds. A mean birth date of 20 June is the first direct estimate of parturition date for UK harbor seals. Information on cumulative births indicated that the peak daily haul‐out count accounted for 77% of total pup production. CMR‐based estimates of lactation duration suggest that 50% of mothers had weaned their pups, when pups were 21‐d old. These results highlight the potential for using photo‐ID techniques to study harbor seal reproductive patterns at sites where intensive capture and marking is not possible for logistic, legislative, or ethical reasons.     

ANNUAL REPRODUCTIVE CYCLE OF THE FEMALE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

Abstract: The annual reproductive cycle is described for the adult female Hawaiian monk seal ( Monachus schauinslandi ) from data collected at Laysan Island (1982–1991) and Lisianski Island (1982–1983) in the Northwestern Hawaiian Islands. Pupping, lactation, weaning, and molting were directly observed, while mating was rarely observed and was, therefore, inferred from the occurrence of mounting injuries and from adult male and female association patterns. Pooled birth rates during the study period were 0.544 for all adult-sized females and 0.675 for females parous in earlier years. For parturient females, pupping peaked in late March and early April, weaning in May, mounting injuries in May and June, and molting in July. For non-parturient females, the median mounting injury and molting dates occurred 17 and 28 days earlier, respectively. Pupping date set the timing of subsequent events in the annual cycle, but the timing of those events was adjusted by loss of the pup or poor physical condition of the female. Individual pupping patterns varied widely. The mean interval for births in consecutive years was 381 days; females that pupped in consecutive years gave birth later each season. Conversely, females who skipped a year or more gave birth earlier their next pupping season.     

THE ROLE OF PREDATION IN THE ECOLOGY OF THE RINGED SEAL IN BARROW STRAIT, NORTHWEST TERRITORIES, CANADA

Predation on ringed seals ( Phoca hispida ) was examined in Barrow Strait between March and May 1984 to 1986. Polar bears were the most important predator. Evidence of bear predation was observed at 18–30% of the ringed seal subnivean structures we located. Ten to 24% of predation attempts were successful, with pups making up 75% to 100% of the seals killed. Bears killed an average of 0.08 to 0.51 seals/km², which comprised 8 to 44% of the estimated annual pup production. Bears were successful on average in 11.3% of their attempts to kill pups hidden inside birth lairs. On southeast Baffin Island where snow was soft and pups were exposed, bears were successful in 33.5% of their attempts to kill a seal. Negative correlations were found between mean snow depth and predation by polar bears ( r = -0.896, P = 0.04, n = 5) in 1985, and between snow depth and the number of predation attempts ( r = -0.613, P = 0.02, n = 14) in 1986.     

Reproductive performance in grey seals: age-related improvement and senescence in a capital breeder

1. Three hypotheses have been advanced to account for age-related improvement in performance: the selection hypothesis predicts improved due to the loss of lower quality phenotypes, the constraint hypothesis predicts individuals improve function, and the restraint hypothesis predicts younger individuals forego or reduce effort because of mortality risks. A decline in age-related performance (i.e. senescence) is predicted by mutation accumulation, antagonistic pleiotropy and disposable soma (wear and tear) hypotheses. 2. Using five measures of performance - birth rate, maternal and pup birth mass, pup weaning mass, weaning success and lactation length - we tested these hypotheses concerning age-related change in reproduction in 279 female grey seals (Halichoerus grypus), ages 4-42 years, over a 23-year period between 1983 and 2005 on Sable Island, Nova Scotia. These females produced 2071 pups. 3. Although body mass of primiparous females increased with age (4-7 years) birth mass of their pups did not, but pup weaning mass did. Second- and third-parity females of the same age as primiparous females gave birth to and weaned heavier pups. However, parity and age were dropped from models when maternal body mass was included. 4. The proportion of females giving birth varied significantly with maternal age, increasing in young females and then declining late in life. Weaning success rate also increased rapidly to about 8 years and subsequently declined in females > 32 years. 5. Generalized additive models indicated nonlinear changes in 3 day body mass (i.e. approximately birth mass) and weaning mass of pups as a function of maternal age, after accounting statistically for the effects of maternal body mass. Mixed-effects, repeated-measures models fitted to longitudinal data further supported the conclusion that pup birth mass and weaning mass vary nonlinearly with maternal age and indicated nonlinear changes in lactation duration. 6. We found some support for the constraint hypothesis, but our findings were not consistent with the selection hypothesis or the restraint hypothesis as the basis for improvement in reproductive performance. 7. Senescence was evident in multiple female and offspring traits, indicating the degeneration in function of several physiological systems as predicted by the disposable soma hypothesis.     

A reproductive study of Weiser-Maples guinea pigs]

The Weiser-Maples (WE) guinea pig strain was introduced by Backshire Co., Ltd. (USA) in 1977. We have been breeding WE strain guinea pigs for skin melanization research. The WE guinea pig colony produced 1271 pups in 417 litters from May 1978 through December 1983. Breeding date are shown below. The mean litter size was 3.05, the stillborn rate was 15.2%, the weaning rate for live-born pups was 93.5% and the sex ratio was 1.01. The average age at first vaginal membrane rupture was 31.4 days at which time body weight was 290.5g. The mean length of the first 7 estrous cycles was about 17 days, with no cyclical variation in length. The mean duration of gestation was 67.9 days. Duration of pregnancy varied with litter size. There was an inverse relationship between litter size and duration of pregnancy. Most of the pups were delivered alive in mid-pregnancy with a parturition range of 56 to 76 days. The probability of pup death depends on gestational length: the lowest incidence of mortality was seen in litters born at 70 days. The mortalities were related to litter size but not to parity. There was an inverse relationship between birth weight and litter size. In WE guinea pigs, the mean weight for a litter of 1 was 120 g; for a litter of 5, the mean body weight was 58g. Male body weights were slightly heavier than female at birth and at weaning age. The mean body weights are shown below, date of birth: female 88.3g, male 93.3g, weaning age (2 weeks): female 181.1g, male 198.8g and 30 weeks: female 758.7g, male 1018.0g. These date for WE guinea pigs are comparable to those of other strains.     

Physiological and behavioral determinants of the aerobic dive limit in Weddell seal (Leptonychotes weddellii) Pups

The aerobic dive limit, as defined by an increase in plasma lactate levels following dives, has to date only been determined in adult and juvenile Weddell seals (Leptonychotes weddellii). However, theoretical aerobic dive limits based on calculated total body oxygen stores, estimated metabolic rates, and dive duration frequencies have been published for several species. Using data collected over the past 3 years in McMurdo Sound. Antarctica, the aerobic dive limit of Weddell seal pups was determined by both the physiological and modeling methods. Time-depth diving recorders deployed on 36 pups between 2 and 14 weeks of age allowed the aerobic dive limit to be predicted from duration-frequency histograms. The aerobic dive limit was also calculated from estimates of total body oxygen stores and predicted diving metabolic rates. Finally, these two estimates were compared with aerobic dive limits determined from post-dive lactate levels in three pups between 5 and 7 weeks old. The aerobic dive limits of pups increased with age, but pup aerobic dive limits were still significantly shorter than those of yearlings and adults. In addition, the aerobic dive limits determined by the three methods were not equivalent for pups, yearlings, or adults, and indicate that care should be taken when modeling methods are used to estimate the aerobic dive limit in other species. Changes in hematocrit, plasma glucose, and plasma lactate levels during and between rest, diving, and recovery in pups were compared to known values for juveniles and adults. Plasma metabolite levels were more highly regulated in older pups, and together with the increasing aerobic dive limit, suggest that Weddell seal pups are not refined divers until after they are weaned, and that their diving ability continues to develop over several years.     

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.     

REDUCED POPULATION GROWTH OF GRAY SEALS AT SABLE ISLAND: EVIDENCE FROM PUP PRODUCTION AND AGE OF PRIMIPARITY

Pup production on Sable Island, Nova Scotia, has been increasing exponentially since the early 1960s and by 1997 Sable Island was the largest gray seal colony worldwide. Using an aerial photographic survey, as in previous years, we estimated pup production in January 2004 to determine if this exponential rate of increase had continued. A total of 33,268 pups was counted on the color positives. When corrected for the proportion pups missed on the imagery (1.106 for the 12th; 1.527 on the 13th), the proportion of pups that died prior to the survey (0.031), and the proportion of pups born before the survey (east colony 0.966, west colony 0.962), estimated total pup production was 41,500 with SE = 4,381. The 2004 estimate indicates that pup production on Sable Island has continued to increase, but suggests that the rate of increase ( r ) may have declined (0.070 compared to previous 0.128). Females from the 1998–2000 cohorts were about 16 times less likely to give birth for the first time at age 4 yr and more than twice as likely at age 6 yr compared to those in the mid-late 1980s. The new estimate of pup production and observed changes in age of primiparity provide the first indication of changes in vital rates of this population. However, additional estimates of pup production and vital rates are needed to confirm this conclusion and to investigate the underlying mechanisms.     

POPULATION SIZE AND BREEDING SEASON OF THE ANTARCTIC FUR SEAL ARCTOCEPHALUS GAZELLA AT HEARD ISLAND—1987/88

Breeding colonies of the antarctic fur seal Arctocephalus gazella on Heard Island (53°10'S, 73°30'E) are situated on the sheltered northern and eastern coasts on flat vegetated terrain near streams and pools. Pupping in the 1987/88 summer began on 21 November, with 90% of births in 26 d. The median birth date was 11 December. Pup counts at Heard Island made in seven breeding seasons from 1962/63 to 1987/88 show an exponential rate of increase of 21%, which may be inflated due to undercounting in early years. The total of 248 births in 1987/88 represents an exponential increase of 37% since the previous year, but pups may have been undercounted then. Based on the number of pups born, the breeding population is estimated at 870–1,120. During the breeding season, the largest number of animals ashore was 835. Many non-breeding fur seals began hauling out from early January and 15,000 animals were estimated to be ashore by late February, a fat larger number than expected from the size of the breeding population. Both the breeding and non-breeding components of the population may be augmented by immigration. The source of immigrants may be undiscovered breeding colonies of this species in the northwestern sector of the Kerguelen Archipelago or the concentration at South Georgia. Further censuses are required at Heard Island to monitor the population growth.     

GROWTH AND DEVELOPMENT IN FREE-RANGING HARBOR SEAL (PHOCA VITULINA) PUPS FROM SOUTHERN BRITISH COLUMBIA, CANADA

Harbor seals ( Phoca vitulina ) are small pinnipeds that are widely distributed throughout the temperate coastal regions of the Atlantic and Pacific oceans. We determined birth mass, neonatal growth rates, weaning age, and weaning mass of NE Pacific harbor seals ( P. v. richardsi ) during a capture-recapture study that spanned the nursing period (Sidney Island, British Columbia, Canada). Of 46 harbor seal pups initially captured, 28 were classified as newborns ( i. e. , < 24 h old). Mean body mass of newborns was 11.2 ± SE 0.31 kg. Pups were individually tagged and recaptured throughout the nursing period. Average daily mass gain during the nursing period was 394 ± 26 g. Mean birth mass of males did not differ significantly from females, although pups found with fetal pelage (lanugo) (21.4% of all newborns) were smaller at birth (9.8 ± 0.44 kg) than non-lanugo pups (11.6 ± 0.33 kg). Mean weaning mass was estimated at 23.6 ± 1.2 kg at a mean weaning age of 32 d ± 1.5 d. While birth and weaning masses differed little from the published data for offshore Sable Island harbor seals ( P. v. concolor ), British Columbia harbor seals are characterized by half the daily mass gain and a longer nursing period.     

DIFFERENTIAL MOVEMENTS BY HARBOR SEAL PUPS IN CONTRASTING ALASKA ENVIRONMENTS

Movement patterns of Alaska harbor seal pups were studied using satellite telemetry during 1997–2000. Mean tracking duration was 277.3 d (SD = 105.8) for Tugidak Island pups ( n = 26) and 171.2 d (108.3) for Prince William Sound (PWS) pups ( n = 27). Movements were similar for males and females and were largely restricted to the continental shelf. Multiple return trips of > 75 km from the natal area and up to ∼3 wk duration were most common, followed by movements restricted to <25 km from the natal area; one way movements from the natal site were rare. Distances moved and home range sizes remained relatively stable or increased gradually from July through winter, then decreased markedly through spring. Monthly movements (maximum distance from tagging location, mean distance from haul-outs to at-sea locations, and home range size) were significantly greater for Tugidak vs . PWS pups. Six of seven pups from each region that traveled farthest and were tracked the longest had returned to their tagging site when their last location was recorded, indicating philopatry or limited dispersal during their first year of life. Seal pups exhibited similar movement patterns in the distinct habitats of the two regions, but differed in the spatial extent of their movements.     

DEVELOPMENT OF DIVING BY HARBOR SEAL PUPS IN TWO REGIONS OF ALASKA: USE OF THE WATER COLUMN

Satellite-linked dive recorders were attached to 53 harbor seal pups in Prince William Sound (PWS) and at Tugidak Island, Alaska, during 1997–1999. We used generalized additive models and bootstrap techniques to describe pup diving behavior during their first year of life. Pups increased their ability to dive during the first 3–6 mo, as indicated by increases in proportion of time in the water (time wet) and maximum dive depth achieved by a pup each day (max-depth) values. Time wet and/or max-depth later decreased, suggesting a seasonal component to diving behavior. Monthly time wet varied from an overall minimum of 0.68 at tagging in July to a maximum of 0.89 in November. Pups spent half of their time wet swimming in water <25 m deep, the shallowest 30% of the available water column. They spent only 5% of their time swimming in the deepest 30% of the available water column, at depths >60–70 m. This strongly suggests they were not feeding on or near bottom during their first year. Average max-depths and deepest actual dives were similar for PWS and Tugidak pups. PWS pups dove deeper sooner and spent less time wet than Tugidak pups during the first few months after tagging, probably as a result of regional bathymetric differences. Diving behavior and body condition suggest that food availability was not likely a major factor in the population decline in PWS during the period of this study.     

Grey seal (Halichoerus grypus) pup production at North Rona: A study of birth and survival statistics collected in 1972

The Grey seal breeding assembly on North Rona was studied throughout autumn 1972. Weekly censuses provided data on birth rate and mortality. It is estimated that 1736 pups were born in 1972. This is about 500 less than the mean production estimate for this assembly during the period 1959–71, possibly a result of disturbance of the breeding site by the expedition. About 600 of the pups died before putting to sea. The mean date of birth and the duration of the pupping season agree closely with earlier work on North Rona, although the mean daily birth rate curve is somewhat irregular, perhaps because there was an unusually dry spell of weather at a critical time in the pupping season. An improved estimate of mortality at this assembly showed a relationship with pup density similar to that observed at the Fame Islands. Techniques for estimating pup production (and consequently total population), though not reliable for determining absolute values, are useful for indicating trends and they suggest a stable population of 8000–9000 animals at North Rona. Such estimates might be improved by the use of better methods of aerial photography.     

THE EFFECT OF BIRTH DATE ON FITNESS OF FEMALE DWARF PERCH,MICROMETRUS MINIMUS (PERCIFORMES: EMBIOTOCIDAE)

Lifetime reproductive success may vary considerably with birth date. I measured phenotypic selection on female birth date in a viviparous teleost fish (Embiotocidae: Micrometrus minimus) by sampling birth-date cohorts over time in Tomales Bay, California. Four episodes of selection were measured: survival from birth to first reproduction, reproductive success in the first breeding season, survival to second reproduction, and reproductive success in the second season. Birth date had a significant impact on fitness in the first two episodes. Early born females were more successful in their first breeding season than late born females (directional selection on birth date), but early born females were less likely to survive the period between birth and first reproduction, relative to females born in the middle of the season (stabilizing selection on birth date). The final two episodes of selection had no detectable effect on birth date. Because of the relationship between birth date and survival in the first year, overall selection on female birth date was stabilizing.     

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.     

Endocrine Changes in Harbor Seal (Phoca vitulina) Pups Undergoing Rehabilitation

Rehabilitating pinniped pups are often admitted to care centers as neonates and generally lack maternal investment and are in poor body condition. Upon admittance to a rehabilitation facility, pups are typically fed a milk replacement formula via gavage, which is switched to frozen fish upon weaning. While rehabilitation has been successful in terms of recovery and release, preweaning growth rates in captivity are consistently lower than in the wild. Indicators of stress (cortisol and total thyroxine; TT4), and standard morphometrics, of harbor seal pups in rehabilitation (n = 20) were determined for both preweaned and weaned pups. Hormone concentrations and standard morphometrics from pups in care were compared with free‐ranging harbor seal pups (n = 59). Pups in rehabilitation gained mass on both milk and fish diets. Preweaned pups had greater mean serum cortisol and similar TT4 concentrations than weaned pups. Free‐ranging harbor seal pups were heavier and longer than preweaned and weaned pups in rehabilitation. The free‐ranging pups had the lowest cortisol and highest TT4 concentrations of any of the pups. These results suggest that weaned pups that have undergone rehabilitation are not physiologically equivalent to free‐ranging weaned pups. Additional research is needed regarding physiological changes in endocrine values during early development under captive care conditions. This information should be useful to marine mammal rehabilitation centers in their development of care protocols and release criteria for rehabilitating harbor seal pups. Zoo Biol. 32:134–141, 2013. © 2012 Wiley Periodicals, Inc.