Microsatellite Analysis of Population Structure in the Endangered Hawaiian Monk Seal

Abstract: The endangered Hawaiian monk seal breeds at six locations in the northwestern Hawaiian Islands. To determine whether significant genetic differentiation exists among these sites, we used microsatellite loci to examine the monk seal population structure at the five largest breeding colonies. Of 27 loci isolated from other seal species, only 3 were polymorphic in an initial screening of one individual from each breeding site. Only two alleles were found at each of these 3 loci in samples of 46–108 individuals. This extremely low variation is consistent with other measures of genetic variability in this species and is probably the result of a recent severe population bottleneck, combined with a long-term history of small population sizes. Although the smallest monk seal subpopulation in this study ( Kure Atoll) showed some evidence of heterozygote deficit, possibly indicative of inbreeding, the next smallest ( Pearl and Hermes Reef) had an apparent excess of heterozygous individuals. Genetic differentiation was detected between the two subpopulations at extreme ends of the range ( Kure and French Frigate Shoals). This trend was significant only at the microsatellite locus for which we had the largest sample size ( Hg6.3: R _ST = 0.206, p = 0.002; allelic goodness of fit G _h = 15.412, p < 0.005). French Frigate Shoals is the source population for translocated animals that have been released primarily at Kure Atoll. Differentiation between these sites consisted of allele frequency differences (with the same allele predominant in each location at all three loci), rather than the preservation of alternative alleles. Although the translocations have had positive demographic effects, we recommend continued genetic monitoring of both the source and recipient populations because translocated individuals are now entering the breeding population.     

Fostering behavior in Hawaiian monk seals: is there a reproductive cost?

Summary 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.     

"Mobbing" in Hawaiian Monk Seals (Monachus schauinslani): The Value of Simulation Modeling in the Absence of Apparently Crucial Data

Several small populations of Hawaiian monk seals ( Monachus schauinslani ) exhibit male-biased adult sex ratios and "mobbing," an aggressive behavior in which adult males injure and often kill adult females and immature seals of both sexes during mating attempts. Mobbing appears to be limiting the growth of some populations of this endangered species. The frequency of mobbing deaths appears to increase as a population's sex ratio becomes increasingly male-biased, although the exact relationship between these two variables (the mobbing response) is unknown. We developed a stochastic demographic model of a small Hawaiian monk seal population using several different assumptions about the mobbing response. We used the model to explore the origins of male-biased sex ratios in monk seal populations and to determine whether it was possible, given the lack of data on the mobbing response, to evaluate the probable effects of alternative management strategies to address the mobbing problem. Small populations (100 to 200 seals) and those with slower growth rates were more likely to develop male-biased adult sex ratios. Almost all of our modeling scenarios supported the immediate removal of males from populations where mobbing occurs. Our conclusions were relatively unaffected when the assumptions regarding the mobbing response were varied. Thus, a model was helpful even when apparently crucial data were unavailable.     

Evidence of a maternal foraging cycle resembling that of otariid seals in a small phocid,the harbor seal

Summary 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.Correspondence to: D.J. Boness     

Shifting Baselines and The Extinction of The Caribbean Monk Seal

The recent extnction of the Caribbean monk seal Monachus tropicalis has been considered an example of a human‐caused extinction in the marine environment, and this species was considered a driver of the changes that have occurred in the structure of Caribbean coral reef ecosystems since colonial times. I searched archaeological records, historical data, and geographic names (used as a proxy of the presence of seals) and evaluated the use and quality of these data to conclude that since prehistoric times the Caribbean monk seal was always rare and vulnerable to human predation. This finding supports the hypothesis that in AD 1500, the Caribbean monk seal persisted as a small fragmented population in which individuals were confined to small keys, banks, or isolated islands in the Gulf of Mexico and the Caribbean Sea. This hypothesis is contrary to the assumption that the species was widespread and abundant historically. The theory that the main driver of monk seal extinction was harvesting for its oil for use in the sugar cane industry of Jamaica during the 18th century is based primarily on anecdotal information and is overemphasized in the literature. An analysis of reported human encounters with this species indicates monk seal harvest was an occasional activity, rather than an ongoing enterprise. Nevertheless, given the rarity of this species and its restricted distribution, even small levels of hunting or specimen collecting must have contributed to its extinction, which was confirmed in the mid‐20th century. Some sources had been overlooked or only partially reviewed, others misinterpreted, and a considerable amount of anecdotal information had been uncritically used. Critical examination of archaeological and historical records is required to infer accurate estimations of the historical abundance of a species. In reconstructing the past to address the shifting baseline syndrome, it is important to avoid selecting evidence to confirm modern prejudices. Puntos de Referencia Cambiantes y la Extinción de la Foca Monje Caribeña     

Does male harassment of females contribute to reproductive synchrony in the grey seal by affecting maternal performance?

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.     

Phylogeography, historical demography, and the role of post-settlement ecology in two Hawaiian damselfish species

Coral reef fish generally have relatively sedentary juvenile and adult phases and a presumed highly dispersive pelagic larval phase, yet previous studies that have tried to relate pelagic larval duration (PLD) to population structure have given inconsistent results. In the present study, the population structures of two damselfishes, Stegastes fasciolatus and Dascyllus albisella, were examined using mitochondrial control region sequences. The two species have similar PLDs (∼25 and 27 days respectively), but consistently differ in their settlement preferences, habitat, and densities in populations throughout the Hawaiian Archipelago, from Hawaii north to Kure Atoll, and south to Johnston Atoll. Information on habitat preferences and population densities were collected between September 2000 and October 2002, and tissue samples for the genetic studies were collected between January and April 2004. Based on the differences in habitat and abundance of the two species, the expectation was that S. fasciolatus would have high genetic variability but little population structure compared to D. albisella, and this was largely confirmed. Stegastes fasciolatus had little population structure in most of the Hawaiian Islands, and D. albisella showed evidence of strong population structure throughout its range. An exception to this pattern was the large difference between the Kure Atoll population of S. fasciolatus and all others. These results suggest that the interaction of several biological factors (e.g. species-specific spawning habitat and season) with environmental factors (e.g. seasonal wind and current patterns) may have more influence on population structure than single life history characteristics, such as the PLD.     

Offspring sex ratio in relation to female size in southern elephant seals,Mirounga leonina

Southern elephant seals Mirounga leonina display extreme sexual dimorphism. In addition females show great variation in size and stored resources at parturition. Therefore they present an excellent opportunity for examination of responses of sex ratio to resource availability. We studied the relationships between the size of southern elephant seal females at parturition and the size and sex of their pups at South Georgia over four breeding seasons. We found a large individual variation in maternal post-partum mass (range 296–977 kg, n=151). Larger mothers gave birth to larger pups, irrespective of the sex of their pup. Male pups were on average 14% larger than females at birth and consequently more costly to bring to parturition. Our results suggest that female southern elephant seals must weigh more than 300 kg if they are to breed at all, and more than 380 kg if they are to give birth to a male pup. Above this threshold the proportion of males among offspring rapidly increases with maternal mass, and stabilizes at a level not significantly different from parity. These results show that smaller females of southern elephant seals vary offspring sex ratio in a way that is consistent with theories on adaptive offspring sex ratio. A smaller mother with a male foetus may benefit from terminating her pregnancy and allocating the resources she saves to her own growth. She could then give birth to and raise a larger pup in the subsequent season.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

High male reproductive success in a low-density Antarctic fur seal (Arctocephalus gazella) breeding colony

Understanding how population density influences mating systems may lead to important insights into the plasticity of breeding behavior, but few natural systems allow for such studies. Antarctic fur seals (Arctocephalus gazella) provide an interesting model system because they breed in colonies of varying densities. Previous studies have largely focused on a high-density site at Bird Island, South Georgia. Here, 13 highly polymorphic microsatellite loci were used to conduct a genetic analysis of a low-density breeding colony of this species at Livingston Island, approximately 1,600 km south of South Georgia. The majority of adults seen ashore (n?=?54) were sampled together with every pup born (n?=?97) over four consecutive years. Paternities were confidently assigned for 34 out of the 97 pups. Two out of 23 sampled males accounted for the paternity of 28 % of all pups sampled during the study and 82 % of the pups with an assigned father. Moreover, a full likelihood pedigree inference method assigned a further eight paternities to a single unsampled male seal that is inferred to have held a territory during the season before the study began. The most successful males in our study easily surpassed the previous record for the total number of pups sired per male seal for the species. Furthermore, we identified two triads of full siblings implying that their parents remated in three consecutive years. These findings suggest that territorial male fur seals may achieve greater success in monopolizing access to breeding females when population density is relatively low.     

Choosing a Management Strategy: Two Structured Decision-Making Methods for Evaluating the Predictions of Stochastic Simulation Models

We describe two structured decision-making methods—one using a hierarchy of goals and a second using ranking on the sum of weighted criteria—that may be useful for many practical conservation problems, particularly when advisory groups evaluate the output of simulation models. We illustrate both methods by applying them to the problem of choosing a management strategy to address the "mobbing" problem in endangered Hawaiian monk seals. Both methods require estimates of the probabilities of various outcomes, such as a population size of more than 400 seals after 20 years under a specific management regime. We used a simulation model of a small monk seal population to generate these probabilities. Both methods provide an explicit, well-documented, and reproducible decision process that helps justify the decision. Furthermore, they are easy for those untrained in decision analysis to understand and use, they focus discussion on management objectives, they facilitate an examination of trade-offs in the light of multiple and sometimes conflicting objectives, they are suitable for use in workshops, and, at least in our example, they lead to management recommendations that are not highly sensitive to minor changes in probability estimates or other factors.     

Male-biased sex ratios in New Zealand fur seal pups relative to environmental variation

A number of models have been proposed to provide adaptive explanations of sex-ratio variation in mammals. Two models have been applied commonly to primates and ungulates with varying success—the Trivers-Willard (TW) hypothesis, and the local resource competition (LRC) hypothesis. For polygynous, sexually dimorphic mammals, where males are larger and disperse more readily, these models predict opposite outcomes of sex-ratio adjustment within the same environmental context (high-resource years: TW—more sons; LRC—more daughters). However, many of the predictions of these two models can vary depending on factors influencing resource availability, such as environmental stochasticity, resource predictability, and population density. The New Zealand fur seal (Arctocephalus forsteri) is a polygynous mammal showing marked sexual dimorphism (larger males), with higher variation in male reproductive success expected. We provide clear evidence of male-biased sex ratios from a large sample of A. forsteri pups captured around South Island, New Zealand during 1996/1998, even after accounting for a sex bias in capture probability. The extent of the bias depended upon year and, in 1998, strong climatic perturbations (El Niño/Southern Oscillation, ENSO) probably reduced food availability. Significant male-biased sex ratios were found in all years; however, there was a significant decline in the male bias in 1998. There was no relationship between sex ratio and population density. We suggest that the sex-ratio bias resulted from the production of relatively more male pups. Under the density-independent scenario, the strong male bias in A. forsteri sex ratios is support for the TW model within an environment of high resource predictability. We suggest that some plasticity in the determination of pup sex among years is a mechanism by which A. forsteri females in New Zealand, and perhaps other otariid seals, can maximise fitness benefits when living in regions of high, yet apparently predictable, environmental variability. We also suggest that much of the inconsistency in the reported sex ratios for otariid seals results from the complex interaction of population density and environmental stochasticity influencing relative food availability over time.     

Breeding site choice fails to explain interannual associations of female grey seals

Consistent, widespread fine-scale preferences for breeding sites may favour the emergence of intrasexual associations and hence sociality. Pairwise association was defined for a terrestrially breeding phocid seal in which intraseasonal colony mixing is limited and mothers spend only 3 weeks ashore breeding. Interannual associations may occur between pairs of breeding female grey seals at North Rona, Scotland because of pupping site fidelity, 90% return rates and limited variability in parturition dates that individual mothers show. Associations can occur when mothers are found together irrespective of location, or because of joint preferences for locations. Mothers were sedentary and were unlikely to interact if they are separated by >20 m. Using spatial and temporal association criteria, 160 of 176 of known mothers had intraseasonal associations with at least one other mother, with 14 of 126 (11.1%) of associations repeated in 2 years. The likelihood of female association in 2 years as a result of site fidelity, parturition date variation and pupping site quality was modelled. Interannual association between mothers that changed their pupping sites by up to the median observed pupping site fidelity (<40 m) was indistinguishable from our models predictions. However, the number of mothers that showed interannual association after displacements of >40 m from their previous years pupping sites was almost five times greater than the model predicted. We conclude that active association between adult female grey seals is not determined solely by habitat preference, and argue that examination of simple sociality in temporarily aggregating species can contribute significantly to socioecological models of social evolution.Communicated by F. Trillmich     

Parental investment in southern elephant seals,Mirounga leonina

Summary The southern elephant seal is among the most sexually dimorphic and polygynous of all mammals: males may be more than 10 times the weight of reproducing females and only the largest 2–3% of males are likely to breed. Current optimization theories of sexual selection predict that evolution would favor greater parental investment in individual males than in females. Because southern elephant seals represent an extreme of polygyny and sexual dimorphism, they might be expected to show a dramatic difference in parental investment in male and female pups. However, in a study of parental investment in elephant seals at South Georgia, using several different methods, we found no such difference after parturition. Mother-pup pairs were immobilized and weighed early in lactation, recaptured near the end of lactation and reweighed. A further 30 pups were weighed an average of five times during lactation to establish the shape of the growth curve and to serve as partial controls for the previous set of animals. Initial post-partum weight in females ranged from 346 to 803 kg (=506, SD=111, n=26). Pup birth weight was related to mothers' post-partum weight in female pups but small females often gave birth to large male pups. Male pups were significantly heavier at birth than females. However, this size difference did not persist. Male and female pups were suckled for the same period, grew at the same rate and were not significantly different in weight at weaning. Mothers lost weight at the same rate regardless of their pup's sex.     

Low Genetic Variability in the Hawaiian Monk Seal

The Hawaiian monk seal (   Monachus schauinslandi) is a critically endangered species that has failed to recover from human exploitation despite decades of protection and ongoing management efforts designed to increase population growth. The seals breed at five principal locations in the northwestern Hawaiian islands, and inter-island migration is limited. Genetic variation in this species is expected to be low due to a recent population bottleneck and probable inbreeding within small subpopulations. To test the hypothesis that small population size and strong site fidelity has led to low within-island genetic variability and significant between-island differentiation, we used two independent approaches to quantify genetic variation both within and among the principal subpopulations. Mitochondrial control region and tRNA gene sequences (359 base pairs) were obtained from 50 seals and revealed very low genetic diversity (0.6% variable sites), with no evidence of subpopulation differentiation. Multilocus DNA fingerprints from 22 individuals also indicated low genetic variation in at least some subpopulations (band-sharing values for "unrelated" seals from the same island ranged from 49 to 73%). This method also provided preliminary evidence of population subdivision (  F'_st estimates of 0.20 and 0.13 for two adjacent island pairs). Translocations of seals among islands may therefore have the potential to relieve local inbreeding and possibly to reduce the total amount of variation preserved in the population. Genetic variation is only one of many factors that determine the ability of an endangered species to recover. Maintenance of existing genetic diversity, however, remains an important priority for conservation programs because of the possibility of increased disease resistance in more variable populations and the chance that inbreeding depression may only be manifest under adverse environmental conditions.     

Continued decline of an Atlantic cod population: how important is gray seal predation?

The continental shelf ecosystem on the Eastern Scotian Shelf (ESS) has experienced drastic changes. Once common top predators are a small fraction of their historical abundance, and much of the current community structure is now dominated by pelagic fishes and invertebrates. Embedded within this food web, Atlantic cod and gray seal populations have recently exhibited nearly opposite trends. Since 1984, cod populations have decreased exponentially at a rate averaging 17% per year, whereas gray seals have continued to increase exponentially at a rate of 12%. We reexamined the impact of gray seals on Atlantic cod dynamics using more than 30 years of data on the population trends of cod and gray seals while incorporating new information on seal diet and seasonal distribution. The closure of the cod fishery over 10 years ago allowed for a better estimation of natural mortality rates. We quantified the impact of seals on ESS cod by (1) estimating trends in seal and cod abundance, (2) estimating the total energy needed for seal growth and maintenance from an energetics model, (3) using estimates of the percentage of cod in the total diet derived from quantitative fatty acid signature analysis (QFASA) and of the size-specific selectivity of cod consumed (derived from otoliths collected from fecal samples), and (4) assuming a gray seal functional response. Uncertainties of the model estimates were calculated using the Hessian approximation of the variance-covariance matrix. Between 1993 and 2000, cod comprised, on average, < 5% of a gray seal's diet. Our model shows that, since the closure of the fishery, gray seals have imposed a significant level of instantaneous mortality (0.21), and along with other unknown sources of natural mortality (0.62), are contributing to the failure of this cod stock to recover.     

Evidence of equal maternal investment in the sexes in the polygynous and sexually dimorphic grey seal (Halichoerus grypus)

In polygynous and sexually dimorphic mammals, parents may be expected to bias their investment towards sons because variation in reproductive success is usually higher among males than among females. Moreover, male reproductive success often depends on adult body size, which, in turn, may depend on the level of parental investment. We therefore predicted that in the grey seal (Halichoerus grypus), a polygynous and sexually dimorphic phocid seal, females should invest more in individual sons than in individual daughters. We found that male pups were born heavier than female pups, but that the growth rates and suckling behaviour were similar for the two sexes. The growth rates and the birth weights were not correlated for the pups of either sex. Mothers did not behave differentially towards offspring of the two sexes, except that mothers of male pups spent more time in visual contact with their pups. Male and female pups had similar activity levels and begged at similar rates. We argue that reports of equal expenditure on the two sexes can be accepted as evidence of equal investment, provided that three assumptions are fulfilled. First, parental care must be costly to the parent. Second, energy expenditure must be the most important component of parental investment. Third, there must be no negative correlation between maternal body condition and the ratio of sons to daughters produced. We argue that these assumptions are met in our study, and that our results provide evidence of equal maternal investment in the sexes in grey seals.     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

Maternal investment in Antarctic fur seals: evidence for equality in the sexes?

Studies of the otariids (fur seals and sea lions), a highly sexually dimorphic group, have provided conflicting evidence of differential maternal expenditure in male and female offspring and, thus, suggestions that they conform to predictions of investment theory are equivocal. Since the mid-1970s, a diversity of research on Antarctic fur seals (Arctocephalus gazella) including studies of their reproductive ecology, lactation energetics, and foraging behaviour have been conducted at Bird Island, South Georgia that have resulted in one of the more complete and diverse data sets for any species of otariid. These long-term data were reviewed to determine whether there was any evidence to support that differential maternal expenditure occurred in Antarctic fur seals. Most of the data examined were collected during five consecutive austral summers from 1988 through 1992 and included years in which local food resources were abundant and scarce. We were unable to detect differences in the sex ratios of pups at birth or sex-biased differences in growth rates estimated from serial data, the number of foraging trips made, the duration of attendance ashore, diving behaviour, suckling behaviour, or milk consumption in any year and in the duration of foraging trips or age at weaning in 2 of 3 years. In addition, we found no evidence of greater reproductive costs between mothers with sons or daughters relative to their reproductive performance the following year. In contrast, sex-biased differences were only found in the duration of foraging trips in 1990, the age at weaning in 1988, and consistently in growth rates estimated from cross-sectional data. We suggest that differential maternal expenditure does not occur in Antarctic fur seals because male pups probably do not gain greater benefit from additional maternal expenditure than female pups. After weaning, males experience a period of rapid juvenile growth over 3–4 years during which time body mass nearly trebles. This growth will almost certainly be dependent upon available food resources then rather than on any maternal expenditure received over the first 4 months of life and, thus, the assumptions of the Trivers and Willard hypothesis are probably invalid for Antarctic fur seals. Received: 10 July 1996 / Accepted after revision: 3 March 1997