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Michael K. Schwartz,a,b Daryl J. Boness,a Catherine M. Schaeff,b Patricia Majluf,c Elizabeth A. Perry,a and Robert C. Fleischera aDepartment of Zoological Research, National Zoological Park, Smithsonian Institution, Washington, DC 20008, USA, bDepartment of Biology, American University, 4400 Massachusetts Avenue, Washington, DC 20016, USA, and cWildlife Conservation Society, Paul de Beaudiez 520, Lima 27, Peru

Our best understanding of marine mammal mating systems comes from land-mating pinnipeds. Logistical problems of observing behavior at sea have limited our ability to make inferences about species with aquatic-mating systems, which comprise over half the pinnipeds. The mating systems of these species likely involve different mating tactics than land-mating species. We used several methods in combination (e.g., animal-borne cameras, radio telemetry, time-depth recorders, and DNA paternity assessment) to provide a comprehensive study of the aquatic-mating tactics of harbor seal males. Males decreased time offshore (26.0 vs 14.8%) and increased time near shore (33.8 vs 43.7%) between premating and mating periods, respectively. Concomitantly, males reduced foraging effort and increased activities associated with competition for females (e.g., visual/vocal displays and threats). As females come into estrus near the end of lactation and spend more time at sea, males reduced their near-shore ranges (4.2 vs 1.0 km2), which were clustered within 1–1.5 km of the beach where females attended their pups. Body mass of males was not a major factor affecting their reproductive behavior. From a small number of paternity assignments to study males, it appears that females select males. These combined results are more consistent with a lek-type mating system than with the territorial or female defense systems characteristic of land-mating pinnipeds.

Summary 1 We studied feeding frequency in free-ranging grey seals using stomach temperature telemetry to test if previously reported sex differences in the diving, movement and diet were reflected in the temporal pattern of foraging success. 2 Data were retrieved from 21 of 32 grey seals from 1999 to 2001, totalling 343 days and 555 feeding events, with individual record length varying from 2 to 40 days (mean: 16·33 ± 2·67 days/seal). 3 Seals fed on 57·8 ± 6·46% of days sampled and had an average of 1·7 ± 0·26 meals per day, but individual variability was apparent in the temporal distribution of feeding as evidenced by high coefficients of variation (coefficient of variation = 69·0%). 4 Bout analysis of non-feeding intervals of six grey seals suggests that feeding intervals of individuals were varied and probably reflect differences in prey availability. Grey seals tended to have many single feeding events with long periods separating each event, as would be expected for a large carnivore with a batch-reactor digestive system. 5 We found significant sex differences in the temporal distribution of feeding. The number of feeding events per day was greater in males (2·2 ± 0·4 vs. 1·0 ± 0·2), as was time associated with feeding per day (56·6 ± 5·8 min vs. 43·9 ± 9·4 min). 6 The number of feeding events varied with time of day with the least number occurring during dawn. Feeding event size differed significantly by time of day, with greater meal sizes during the dawn and the smallest meals during the night. 7 The length of time between meals increased with the size of the previous meal, and was significantly less in males (541·4 ± 63·5 min) than in females (1092·6 ± 169·9 min). 8 These results provide new insight into the basis of sex differences in diving and diet in this large size-dimorphic marine predator.

We studied female and male parental roles in two sympatric auks, thick-billed murre (Uria lomvia) and razorbill (Alca torda), with initial biparental care at the breeding site and later exclusively paternal care at sea. Our study addressed the following two questions: Why do males accompany chicks to sea?, and How do the sexes allocate parental effort at the breeding site before parental care at sea begins? We tested the hypothesis that males care for chicks at sea because they are in better condition at the time of chick departure as a result of femalebiased parental effort at the breeding site (‘nest’). Breeding success and duration of chickrearing did not differ between the two study years in either species at the Gannet Islands, Labrador. At the breeding colonies, females of both species provided more food (murres: 2.84 ± 0.18 loads day−1; razorbills: 2.02 ± 0.11 loads day−1) to their chicks than males (murres: 2.26±0.12 loads day−1; razorbills: 1.42±0.09 loads day−1), and males spent more time brooding the chicks. These differences were chick-age dependent in both species, with females providing more meals to chicks older than two weeks. Razorbill males spent more time with chicks greater than two weeks old, while murre male’s attentiveness of brooding did not vary with chick age. In both species, males (murres: 3.04 ± 0.3 h day−1; razorbill: 3.30±0.2 h day−1) performed longer foraging trips with meal delivery than females (murres: 1.23 ± 0.4 h/day; razorbill: 2.50 ± 0.4 h day−1). Thick-billed murres showed a consistent diurnal pattern of egg and chick attendance: females were usually found at the breeding site during the day whereas males were found there early in the morning and at night. In contrast, razorbill’s timing of attendance was much more variable and did not differ between sexes. Despite these differences in timing of breeding site attendance between species, males of both species spent twice as much time as females engaged in the defence of the egg or chick at the breeding site, which suggest male-biased capability of protecting the chick at departure. Overall our data indicated different female and male parental roles at the breeding site but not a female-biased allocation of time, energy and risk as predicted. In fact, males seem to provide equal if not more parental effort than females prior to the time of colony departure. We propose that the patterns of parental roles found between sexes is the result of a chain of events favouring male involvement in chick brooding and care at sea.

The purpose of this study was to examine the importance of body size and body composition as determinants of conditional mating tactics exhibited in male grey seals. We combined behavioral observations with measures of energy expenditure and success on 42 known-age individuals during the breeding seasons of 1997–2001 at Sable Island, Canada. Males with a large body mass arrived at the breeding grounds with more body fat and body energy and catabolized less body protein than smaller males. Males consumed 1.9 6 0.2 MJ day � 1 , and those with a smaller percentage of body fat had higher rates of food energy intake. The amount of body energy on arrival was positively correlated with the duration of the breeding period. Males that exhibited the primary mating tactic of consortship were heavier, had absolutely more body fat and body energy, and sustained breeding longer than those males that did not exhibit the primary tactic. Amongst all males, body mass showed a quadratic relationship with the number of female consorts mated and the estimated number of pups sired. Thus, intermediate-sized males mated with the most consorts and achieved the highest success. Intermediate body size may be optimal during breeding due to greater agility in male combat. Body size was an important determinant of mating tactics used by male grey seals. A large body size provided an energetic advantage of greater endurance while an intermediate body size may provide greater competitive ability in acquiring consortships. Key words: body size, conditional mating tactics, endurance rivalry, energetics, Halichoerus grypus, pinnipeds, reproductive behavior. [Behav Ecol]

Predators face decisions about which prey to include in their diet in order to maximize fitness. The foraging tactics used to capture prey and the resulting profitability of prey influence these decisions. We present the first evidence of prey-dependent foraging tactics and prey profitabil- ity in a free-ranging pinniped. We studied 39 adult male harbour seals Phoca vitulina at Sable Island, Nova Scotia using an animal-borne video system. Each male wore the camera system for 3 d during which 10 min video samples were recorded every 45 min from 06:00 h, resulting in approximately 3 h of videotape per male and a total of 1094 capture attempts of identified prey. Males foraged mainly on sand lance Ammodytes dubius and flounders (Pleuronectids), but salmonid and gadoid fishes were occasionally pursued. Foraging tactics differed among and within prey types based on differ- ences in prey behaviour. Sand lance was both a cryptic prey, when in the bottom substrate, and a con- spicuous schooling prey. Seal swimming speed, handling time and capture success differed between cryptic and conspicuous sand lance. The highest capture success and handling time was recorded for flounders. Estimated profitability, i.e. net energy intake per unit time, also differed with prey type and prey size. Our results suggest that diet selection may have important implications on the foraging energetics of pinnipeds.

We analyzed patterns of egg laying and breeding frequency of Humboldt Penguins (Spheniscus humboldti) between 1993 and 1997 at Punta San Juan, Peru. Egg-laying extended from mid-March to the first week of December, showing two well-defined peaks in April and August–September. The extended breeding period of these birds was the result of individuals having a second clutch. About half of the females (n =189) had two clutches per year, most of which were double broods (73%). The date of completion and outcome of reproduction, or whether a change of mates occurred from the previous year, did not affect timing of egg laying. The majority of first clutches (62%) were laid in April each year. Two-clutch breeders that started laying eggs early in April had a higher breeding success than those starting in late April, and double brooders had greater success than single brooders. Two-clutch breeders started to lay eggs earlier than single-clutch breeders. Taking into account that a penguin breeding cycle (from egg laying to fledging) lasts ∼4 months, laying eggs early in April increases the chance of rearing two successful broods per year. During three consecutive years, females tended to have two clutches instead of only one clutch and an average breeding success of 4.54 fledglings over 3 years. Having as many clutches as possible when conditions are favorable appears to be a strategy used by Humboldt Penguins to maximize their lifetime reproductive success within a productive but unpredictable environment.

Marine animals equipped with data recording devices sometimes show changes in their foraging behaviour, which can indicate an adverse effect of the instrument. We attached time-depth recorders (TDRs) and radio-transmitters to study foraging behaviour in Humboldt Penguins Spheniscus humboldti and observed an increase in the duration of foraging trips. We suggest that further tagging of this endangered species be approached with caution since this increase in trip duration may negatively affect reproductive and foraging success.

Summary 1. Female mammals increase energy expenditure during lactation to support the high cost of milk production. The extent to which lactation in a small phocid species, the Harbour Seal Phoca vitulina L., was fuelled by food vs body stores, how this allocation varied with maternal body mass and the consequences of maternal expenditure on offspring growth were studied. 2. The proportional body composition of 30 females was independent of initial postpartum body mass, but larger females had absolutely more stored energy than smaller ones. 3. Females lost 32% of postpartum body mass and 62% of body energy by late lactation; 97% of energy loss was derived from body fat. Percentage loss of body energy was independent of initial body mass, indicating that females limit their allocation of body stores to offspring by expending a constant proportion of stores rather than a constant amount. 4. Females spent more time diving and individual dives were deeper and longer as lactation progressed. By late lactation, these characteristics of diving were inversely proportional with initial postpartum mass. 5. During early lactation, female expenditures were covered mainly by a reduction in body energy stores. By late lactation, food intake increased six-fold but the extent of this increase varied inversely with postpartum mass. 6. Pup growth rate and weaning mass were positively related to postpartum mass and total daily energy expenditure of females, but were independent of the source of energy used by females during lactation. Pups of heavy females had higher survival than pups of light females. 7. Our results support the hypothesis that maternal body mass is an important determinant of lactation strategies in pinnipeds.

During observations at a breeding colony of Humboldt Penguins (Spheniscus humboldti), we noted five male penguins enter nests occupied by breeding pairs and engage in aggressive interactions with the resident adult. These nest intrusions resulted in egg loss or infanticide, and accounted for 11.1% of breeding failure in the colony. Two intruder males ultimately mated with respective resident females. This behavior had not been observed previously in Humboldt Penguins and we suggest that it may occur when males are unable to obtain mates through advertising at the nest. Intrusion behavior may slow down population recovery in this endangered species.

We studied effects of maternal age, body mass, and parturition date on birth mass and stage of development at birth in harbor seals (Phoca vitulina) on Sable Island, Nova Scotia, over a 10-year period. As predicted, effects of maternal mass and age on pup traits varied with maternal age. Premature pups, characterized by extensive fetal pelage termed lanugo, weighed 20% less than newborns without lanugo and were born early in the season to lowbody-mass females 4‐6 years of age. Among intermediate-age females (7‐10 years old), percentage of lanugo on pups was correlated negatively with female postpartum body mass but not with female age. Thus, maternal condition rather than age had stronger effects on stage of development of offspring at birth. Overall, maternal age explained 54% of the variance in birth mass, whereas postpartum mass of females explained only 20% of the variance. These relationships did not differ with sex of the pup, although male pups were 3.7% heavier than female pups. Effects of maternal age were strongest among young females; however, reproductive parity was more important than age itself in determining offspring birth mass.

When closely related species breed in sympatry, and where hybrids have lower fitness, reinforcement theory predicts that selection should favour mechanisms that reduce the probability of interspecific matings. If this situation arises among species that exhibit resource defence polygyny where males and females of different species reside in the same territories, there may be some conflict between mate choice based on territory-holding ability (sexual selection) and mate choice for correct species. We investigated this in a population of fur seals where three species are sympatric and where some females breed in the territories of heterospecific males, and where interspecific matings and hybrid pups are observed. The territorial status of males and the birthing sites of females were determined during daily observations, as were the movements of males and females, the location of matings and mating partners. DNA extracted from skin samples was used to determine paternities using DNA fingerprinting and the mtDNA genotype of individuals. Individuals were also classed on the basis of species-typical phenotype. We found that extra-territory inseminations (ETIs) were significantly more prevalent (67%) when territorial males and resident females were of different phenotype than when of similar phenotype (27%), but mtDNA genotype had no effect on the rate of ETIs. ETIs were probably by males with the same phenotype, as pups born to these females in the following season had the same phenotype as their mothers, suggesting they were not hybrids. These results suggest that within the resource defence polygynous mating system of these sympatric fur seals, female mate choice is more influenced by male phenotype than genotype. Contrary to our predictions, our study indicates that potential conflict between mate choice based on sexual selection and species recognition is unlikely, because females have some capacity to discriminate between males both within and between species on phenotypic traits additional to those under sexual selection. Although at least 25% of the pups born in this study were hybrid, this study can only support reinforcement theory if hybrids have reduced fitness. The fitness of hybrids among the species studied is currently unknown.

The energy expenditure of breeding male harbour seals, Phoca vitulina, on Sable Island, Nova Scotia, was investigated by measuring changes in body mass, body composition, and water flux using isotope dilution. Seals lost 0.47% +/- 0.04% (n = 34) of their initial mass per day during the breeding season (4 wk), and fat, water, and protein accounted for 64.3% +/- 4.8%, 27.8% +/- 3.3%, and 6.9% +/- 1.4% of this mass loss, respectively (n = 31). Total energy expenditure was estimated as 33.3 +/- 1.9 MJ d(-1), or 3.9 +/- 0.2 W kg(-1) (n = 17), similar to rates measured in terrestrially mating pinniped species. However, unlike terrestrially mating pinnipeds, male harbour seals did not fast during the breeding season, and energy intake from foraging accounted for 61.8% +/- 4.0% of the total energy expended. Males derived most of their expended energy from food intake early in the breeding season. However, as oestrus females became increasingly available, reduced rates of food intake in males were coupled with increased rates of total energy expenditure. Larger males expended significantly more energy from body stores and more total energy than smaller males. Male harbour seals appeared to balance the energetic costs of reproduction against the constraints of small body size by foraging during deep-diving trips before the appearance of oestrus females and by opportunistic feeding throughout the breeding season while at sea. We suggest that size dimorphism may be less pronounced in aquatically mating pinnipeds partly because the temporal and spatial separation of foraging and reproduction is less distinct than it is for terrestrially breeding pinnipeds.

Species of the suborder Pinnipedia belong to three families (Phocidae-true seals; Otariidae­ fur seals and sea lions; and Odo­ benidae-walruses) and are distinc­ tive ~mong mammals in that they are neither exclusively land dwelling nor excl,usively ocean dwelling. Pinni­ peds spend the majority of their time at sea foraging; however, with the exception of the walrus, they appear to require land (or sea ice) on which to give birth and nurse their young (Bonner 1984, Oftedal et a1. 1987a). Consequently, nursing of young and foraging for food must be spatially and temporally separated. Although pinniped, females and males aggre­ gate during the breeding season, fe­ male pinnipeds, unlike many other carnivores, are the sole providers of nutritional resources to the young; there is no male assistance or ex­ tended social group that assists the

Harp seals (Phoca groenlandica) give birth on pack ice and nurse their pups for about 13 d. We hypothesized that lactating females would transfer milk and energy to pups at a high rate to compensate for the brevity of lactation. To test this hypothesis, the milk intake of five pups was measured over a 6-8-d period after administration of deuterium oxide; the initial age of pups was estimated to range from 0 to 5 d postpartum. Body water content of 17 pups was also measured by isotope dilution, and body composition of 12 pups was determined by chemical analysis. The proximate composition of harp seal milk (n = 21) was also assayed. Harp seal pups contained little fat (3%) at birth, but 61% of the increase in body mass (average 2.3 kg/d) was fat, with the result that fat accounted for 47% of body mass at weaning. Body water estimated by isotope dilution was not significantly different from that measured directly (n = 9 paired samples); water composed 72% of lean body mass. From birth to weaning, milk lipid ...

Proximal mechanisms underlying a faster growth rate in male compared to female California sea lion pups were investigated. Males are significantly larger at birth than females. Specifically, we asked if differential maternal investment contributed to enhanced male growth via: (1) larger mothers having disproportionately more male pups, (2) more time and energy put into foraging by mothers of male pups, and (3) greater milk production in mothers of male pups. We also considered four aspects of differential energy utilization and acquisition by male and female pups: (1) male pups attempting to save energy for growth by changes in behavior, (2) longer suckling bouts with mother and more sneak suckling of non-filial females by male pups, (3) lower maintenance costs in males via a lowered resting metabolic rate, and (4) increased assimilation efficiency in males. Our study showed that there are no differences in the size of females or length of foraging trips for mothers of male and female pups. Male pups received more milk from their mothers, but the difference was no longer significant when the larger body size of males was considered. There were no differences in either the activity budgets or suckling behavior of male and female pups. Male pups, however, did have lower resting metabolic rates than females. We conclude that enhanced male perinatal growth is a consequence of a larger size at birth, proportionally more milk from mothers to support the greater demands of larger body size, and lower maintenance costs due to a lower resting metabolic rate.

We investigated the possibility that male harassment of lactating females differed in relation to time of birth in the grey seal, Halichoerus grypus, on Sable Island, Nova Scotia. This was done by comparing the frequency of male disturbances, maternal performance and pup growth for females that either gave birth during the peak of the pupping season or after the peak. Of the females, 58% gave birth in a 7-day period near the beginning of the pupping period, when the operational sex ratio was 2–4 females per male. Late in the pupping period the operational sex ratio reversed to about 1 female for every 2 males. The relative frequency of disturbances by males was significantly greater for late-pupping mothers than for peak-pupping ones (1.9 vs. 1.4 encounters/h). Females that gave birth late also were disturbed by males 3 times more often than females that gave birth during the peak (3.4 vs. 1.1 % of observation time). Late-pupping mothers spent 22% less time suckling (4.0 vs. 5.1 % of observation time), had 30% slower growing pups (1.7 vs. 2.4 kg/d), and weaned pups that were 16% lighter (45.6 vs. 54.0 kg). The effect of birth time on pup mass gain and weaning mass was not attributable to factors such as maternal mass, pup birth mass or pup sex. We conclude that the reduced maternal performance is likely the result of the increased male harassment. As reduced weaning mass can lead to reduced juvenile survival, male harassment may have contributed to the enhanced reproductive synchrony in this species.

Lactation strategies in the two largest families of seals have been characterized as a phylogenetic dichotomy, with sea lions and fur seals (Otariidae) exhibiting foraging cycles and true seals (Phocidae) a strategy of fasting. We show that a lactating phocid, the harbor seal, Phoca vitulina, has a foraging cycle similar to that of otariids. Time-depth recorders attached to lactating harbor seal mothers revealed that 9 of 11 females began bouts of diving, averaging 12–40 m, by mid-lactation (12 days). During the remainder of lactation, females made an average of seven diving trips, lasting about 7 h. They returned to the rookery during the interval between successive bouts to nurse their pups. Diving was more frequent during daylight than at night and diving bouts increased in duration as lactation progressed. The diving behavior of females that had weaned their pups and previously collected data from stomach lavage, suggest that the bouts of diving represent successful foraging. We propose that the lactation strategy of the harbor seal is intermediate to that of the otariids and other phocids studied. The harbor seal has a foraging cycle like the otariids, but typically resembles other phocids in length of lactation, rate of mass gain in pups, and in milk fat content. As harbor seals are among the smallest phocids, and only slightly larger than most otariids, it seems likely that maternal size constrains the amount of stored energy harbor seal females can bring to the rookery, forcing them to start feeding during the lactation period.

Hooded seals (Cystophora cristata) have a lactation period of 4 d. The rate of milk energy transfer must be very high to ensure that pups are weaned with sufficient energy reserves to withstand the lengthy postweaning fast. Milk energy transfer was measured by isotope dilution (n = 6), and energy deposition was assayed by chemical analysis of pups at birth (n = 4) and weaning (n = 5). Estimates of body water content obtained by isotope dilution and direct analysis were not significantly different. Water constituted only 71.7% ± 0.50% (SE) of lean body mass at birth, indicating developmental maturity. Newborn pups were also high in fat (14.0% ± 0.78%) and energy (10.1 ± 0.25 MJ/kg). Pups consumed 7.52 ± 0.469 kg/d milk containing 187 ± 11.6 MJ/d. Total milk and energy yields were estimated as 30.1 kg and 746 MJ. Pups deposited 88% of ingested fat, 41% of protein, and 84% of energy. Of the weight gain, 82% was fat. The energy content of pups increased nearly fourfold from birth to weaning, but protein conte...

In phocid species in which females fast throughout lactation, maternal energy reserves at parturition must support lactation. Thus, differences in body size and energy stores may influence the magn...

Abstract Recent studies of fur seals and sea lions (Otariids) which are cold water adapted, indicate that social behaviour in hotter climes is constrained by the form and availability of cool substrate provided by the rookery environment. This first study of the behaviour of the Juan Fernández fur seal provides new evidence of this relationship and the extent to which thermoregultory requirements can affect social behaviour in pinnipeds. Females of this species on Alejandro Selkirk Island made daily movements from inland pupping and rest sites to the shoreline and into the water in response to rapid increases in solar radiation to levels exceeding 1.3 cal/cm2/min. Thirty percent of these females floated and groomed offshore in the afternoon in areas protected from the surf by offshore islets and rocky reefs. Males held territories on land either along the shoreline (39%) or in land-locked areas (45%) as is typical of fur seals, or held completely aquatic territories (16%) that encompassed the site where females floated. The average aquatic male, present on territory primarily during the afternoon hours when females floated offshore, achieved as many copulations as did the average land-locked or shoreline male. Land-locked males often abandoned their territories for short periods (45 minutes on average) in response to increasing solar radiation in the afternoon and at a time when female numbers on land were low. The existence of aquatic territoriality as a successful mating strategy has not yet been documented for any other otariid. The occurrence of this behavioural strategy is likely a product of the interaction of thermoregulatory constraints and topographical features which together promote predictable aggregations of females offshore. This study further substantiates that constraints on female aggregation appear to dictate the range and mode of male mating strategies in otariids.

Gray seals (Halichoerus grypus) undergo a terrestrial postweaning fast (PWF) that depletes energy reserves acquired during the suckling interval. Plasticity in PWF duration may ensure that pups of variable body condition depart for sea with adequate energy reserves. To test this hypothesis, we examined body condition of 30 gray seal pups at weaning and monitored their PWF duration. On average, fat accounted for 47.3% +/- 0.7% of their 53.2 +/- 1.3-kg weaning mass. Although fasting duration averaged 21 +/- 1.1 d (n = p28), there was considerable variation in fasting duration (9 to > 31 d) and the resulting age when pups departed to sea (26 to > 49 d). Percent fat at weaning(38.6%-54.6%) was positively correlated with fasting duration(n = 28, r = 0.376, P = 0.0489). In contrast, total body gross energy (735.3-1,447.4 MJ) and body mass (39.0-66.0 kg) were not correlated with fasting duration. Thus, body composition,not overall body reserves, predicted fasting duration, but the effect was weak, indicating that other factors also account for the observed variation in fasting duration. We speculate that pups with greater percent fat more effectively utilized lipid and conserved protein while meeting metabolic costs throughout the PWF. As a result, fatter pups extended the PWF duration,which may be critical for development of diving physiology and may have facilitated their survivorship to age 1.

Fostering behavior has been reported in a large number of mammal and bird species although the relative frequency of its occurrence in most species is unknown. A commonly held view is that fostering is costly to the parent(s) engaged in it. However, empirical studies of fostering are few, and measures of either cost or benefit are even rarer. During a study of individually marked Hawaiian monk seal mothers and pups, observed over the course of maternal care, I found that 87% of 30 females fostered pups. Females sequentially fostered an average of 2.3 pups (range: 1–5 pups) during the approximately 40-day lactation. The median proportion of lactation spent fostering was 34% (range: 5%–90%). Confusion during aggressive interactions appeared to be the major antecedent of fostering and may be understandable in terms of the spatial pattern among females. The density of females with pups was relatively low for a land-breeding seal (1.5 females per 1000 m2), and the typical spatial pattern indicated a tendency toward dispersion. Yet, movements of females and pups to and from water occasionally leave females within a meter or two of each other. Several measures of the immediate reproductive cost of fostering were obtained, including: the length of time suckled by pups, the size of pups at the end of suckling, and survivorship to 1 year of age. There was no correlation between these measures for individual pups and the extent to which their mothers fostered, indicating that the high levels of fostering may be maintained in monk seals because they are not selected against.

Summary 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. ∼ 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.

To successfully transition from nursing to foraging, phocid seal pups must develop adequate diving physiology within the limited time between birth and their first independent foraging trip to sea. We studied the postpartum development of oxygen stores in gray seals (Halichoerus grypus, n=40) to better understand the ontogeny of diving capacity in phocids. Hemoglobin (Hb), hematocrit (Hct), blood volume (BV), and myoglobin (Mb) levels in newborn (3 d postpartum [DPP]) and newly weaned (17+/-0.4 DPP) pups were among the lowest measured across age classes. During the pups' terrestrial postweaning fast (PWF), Hb, Hct, mass-specific BV, and Mb increased by 28%, 21%, 13%, and 29%, respectively, resulting in a 35% increase in total body mass-specific oxygen stores and a 23% increase in calculated aerobic dive limit (CADL). Although Hb and Hct levels at the end of the PWF were nearly identical to those of yearlings, total body mass-specific oxygen stores and CADL of weaned pups departing for sea were only 66%-67% and 32%-62%, respectively, of those for yearlings and adult females. The PWF represents an integral component of the physiological development of diving capacity in phocids; however, newly independent phocids still appear to have limited diving capabilities at the onset of foraging.

The Concorde produces audible sonic booms as it passes 15 km north of Sable Island, Nova Scotia, where gray and harbor seals occur year round. The purpose of this research was to assess how sonic booms affect these seals. The intensity of the booms was measured and three types of data (beach counts, frequency of behavior, and heart rate) were collected before and after booms during the breeding seasons of the two species. In addition to the data taken during breeding, beach counts were made before and after booms during the gray seal moult. The greatest range in overpressure within a single boom was 2.70 psf during gray seal breeding and 2.07 psf during harbor seal breeding. No significant differences were found in the behavior or beach counts of gray seals following sonic booms, regardless of the season. Beach counts and most behaviors of harbor seals also did not differ significantly following booms, however, harbor seals became more vigilant. The heart rates of four gray seal mothers and three pups showed no clear change as a result of booms, but six male harbor seals showed a nonsignificant tendency toward elevated heart rates during the 15-s interval of the boom. These results suggest sonic booms produced by the Concorde, in level flight at altitude and producing on average a sonic boom of 0.9 psf, do not substantially affect the breeding behavior of gray or harbor seals.

Researchers were on Sable Island, Canada (440°N, 600°W), for the 1997 grey seal (Halichoerus grypus) breeding season to study the affects of sonic booms on seal behavior. The physical parameters of 36 sonic booms associated with the Concorde’s trans‐Atlantic flights were recorded. The average minimum overpressure was −0.568 PSF and mean maximum overpressure was 0.935 PSF. Data were collected on frequency of aggression, movements, and distances moved in 10‐min periods before and after booms to detect changes in general agitation of pups and adults. The number of mother‐pup pairs nursing and number of nursing bouts interrupted during the minute of sonic booms were compared to those in randomly chosen minutes to measure immediate startle responses of seals. There was no difference in the behavior of adults nor pups between 10‐min periods before and after booms, nor did their behavior differ in the minute of booms when compared to randomly chosen control minutes. Female heart rates did not vary before, during and after booms but pup heart rates were variable. A similar study on Sable Island harbour seals (Phoca vitulina) is being conducted for comparison.

Breeding activity budgets of male gray seals were estimated to determine how they are fine tuned to changing demands and to determine whether or not the limited body reserves produced by fasting act to minimize energetically demanding activities. Behaviors were divided into nonsocial and social activities with each category including both active and inactive behaviors. The overall frequency of social behavior did not change throughout a male's tenure but aggressive behavior decreased while sexual behavior increased. Even though resting accounted for the greatest proportion of activity, males that were most active had the greatest copulatory success, suggesting that fasting does not place severe limitations on their activity levels.

Observed changes in maternal investment due to an environmentally induced decrease in food supply (the 1983 El Nino-Southern Oscillation) are compared witha priori predictions for the California sea lion (Zalophus californianus). Changes in behavior, growth and mortality of off-spring were also examined. Data collected in the first two months postpartum for the years before (PRE), during (EN), and the two years after (POST1 and POST2) the 1983 El Nino indicate that females initiated postpartum feeding trips earlier during the food shortage, and spent more time away on individual feeding trips in both the El Nino year and the year after. Perinatal sex ratios (♀:♂) in the years PRE, EN, POST1 and POST2 were 1:1, 1.4:1, 1.1:1 and 1:1.4, respectively. Fewer copulations were observed during the El Nino year, but this difference was not statistically significant. Pups spent less time suckling in the food shortage year and the year following, but attempted to sneak suckle more. Pups were less active and played on land less in the El Nino and following year. Finally, maternal investment as measured by milk intake of offspring was decreased, pups grew more slowly, and suffered increased mortality during the food shortage year. Despite expected sex differences in maternal investment and pup behavior in response to food shortage, there were no sex-biased differences in response in either females or pups. As expected, the food shortage did not affect adult males since they migrate north during the non-breeding season where the environmental perturbation was less severe.

Resting metabolic rate and body temperature of California sea lion pups (Zalophus californianus: Otariidae) were measured for 39 animals (12,000 g, mean body mass) captured from breeding areas on San Nicolas Island, California. These 3-5-wk-old pups had a mean specific resting metabolic rate of 0.780 ± .010 ml O₂ g⁻¹ h⁻¹, which is 2.380 times the value predicted by Kleiber for adult mammals with similar body mass; mean body temperature was 37.8 ± .08 C. This level of metabolism is similar to those reported for young seals (Phocidae). The heat-transfer coefficient (h) was 24.6 times that predicted from body mass, and it appeared that some individuals were on the verge of hypothermia at ambient air temperatures below 19 C. Although high metabolic rates are typically associated with high growth rates in terrestrial mammals, high heat loss associated with relatively poor insulation in sea lion pups may require allocation of relatively large amounts of energy to thermoregulation (=maintenance), thus limiting t...

Isotope dilution was used to measure milk intake of 43 California sea lion pups on San Nicolas Island, California, from 1982 to 1984. Deuterium concentration was monitored in serial blood samples collected at about 5-day intervals for 3 wk after isotope administration. Estimated milk intakes differed significantly between male (723 ± 31.0 g ․ day⁻¹) and female (609 ± 24.0 g ․ day⁻¹) pups but did not increase from the first to second month postpartum. On a metabolic size basis (weight0.83), gross energy intakes did not differ between male and female pups but did differ between years, apparently as a consequence of the 1983 El Nino. The mean energy intakes (kcal ․ day⁻¹) of sea lion pups in the first (61 · W0.83) and second (308 · W0.83) months were high relative to terrestrial species. Regression of energy intake on growth rate predicted that maintenance energy requirements (kcal ․ day⁻¹) were also high (168 and 252 · W0.83 in the first and second months). The fact that California sea lion pups devote a la...