The Behavioral Ecology of Maternal Effort in Fur Seals and Sea Lions

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Abstract
Otariid seals form a group of similar species distributed from subpolar to tropical seas. Many species were recently studied under conditions of low and high food abundance during major oceanographic disturbances. Evidence for the phenotypic flexibility of and constraints on the rearing strategy of these species derives from inter- and intra-specific comparisons and experimental measurements. I review how changes in food abundance influence foraging behavior and energetics of mothers, pup growth rates, milk composition, weaning age, and the fertility cost of pup rearing. Eared seal females rear young by alternating between lactation ashore and foraging at sea. Subpolar fur seals always wean pups at four month of age, a trait which seems to be genetically fixed and adaptive in their highly seasonal environment. Temperate and tropical fur seals and sea lions can respond to changes in food abundnace by increasing or decreasing time to weaning. During pup rearing, mothers regulate body mass to different absolute values when abundance of food resources changes. This seems to be a constraint caused by reduced foraging efficiency at low food availability. Most species feed exclusively during the night, but sea lions (Zalophus californianus) and Northern fur seals (Callorhinus ursinus) also forage during the day by diving deeper. Females of some species increase the rate of at-sea energy expenditure at low food abundance while keeping the duration of foraging trips constant. Females of other species stay at sea longer working at the same rate. The latter species may be food specialist with a narrower time window for efficient foraging. Pups grow slower when mothers increase time at sea, and at very low food abundance pups may starve. Pups have little influence on the duration of maternal foraging trips, but may diminish the mothers' time ashore by increasing the rate of milk extraction and prolonged non-nutritive sucking. The impact of long maternal at-sea times is reduced by increasing milk fat content as time at sea increases. This mechanism does not fully compensate pup energy intake rate for longer trip durations. Non-migratory species can partly compensate for reduced pup growth rates by lengthening the lactation period. In the Galapagos fur seal (A. galapagoensis), this entails a considerable cost to the mother by reducing her future fertility. Lactation reduces the probability of a successful simultaneous pregnancy and, if pregnancy succeeds, sibling competition for maternal milk ensues. Competition is usually won by the older sibling leading in many cases to the death of the newborn. The rate of energy transfer to pups is high in food-rich years and low in scarce ones. Because pups are more likely to be weaned as yearlings when juvenile growth rate is high than when it is low, high energy transfer to the pup in year "a" reduces the cost of reproduction incurred in year "a+1". The flexibility of the rearing strategy of temperate and tropical species permits mothers to adjust phenotypically to variance in food availability thus partly masking the theoretically expected trade-offs in the life history of these species. Otariid seals form a group of similar species distributed from subpolar to tropical seas. Many species were recently studied under conditions of low and high food abundance during major oceanographic disturbances. Evidence for the phenotypic flexibility of and constraints on the rearing strategy of these species derives from inter- and intra-specific comparisons and experimental measurements. I review how changes in food abundance influence foraging behavior and energetics of mothers, pup growth rates, milk composition, weaning age, and the fertility cost of pup rearing. Eared seal females rear young by alternating between lactation ashore and foraging at sea. Subpolar fur seals always wean pups at four month of age, a trait which seems to be genetically fixed and adaptive in their highly seasonal environment. Temperate and tropical fur seals and sea lions can respond to changes in food abundnace by increasing or decreasing time to weaning. During pup rearing, mothers regulate body mass to different absolute values when abundance of food resources changes. This seems to be a constraint caused by reduced foraging efficiency at low food availability. Most species feed exclusively during the night, but sea lions (Zalophus californianus) and Northern fur seals (Callorhinus ursinus) also forage during the day by diving deeper. Females of some species increase the rate of at-sea energy expenditure at low food abundance while keeping the duration of foraging trips constant. Females of other species stay at sea longer working at the same rate. The latter species may be food specialist with a narrower time window for efficient foraging. Pups grow slower when mothers increase time at sea, and at very low food abundance pups may starve. Pups have little influence on the duration of maternal foraging trips, but may diminish the mothers' time ashore by increasing the rate of milk extraction and prolonged non-nutritive sucking. The impact of long maternal at-sea times is reduced by increasing milk fat content as time at sea increases. This mechanism does not fully compensate pup energy intake rate for longer trip durations. Non-migratory species can partly compensate for reduced pup growth rates by lengthening the lactation period. In the Galapagos fur seal (A. galapagoensis), this entails a considerable cost to the mother by reducing her future fertility. Lactation reduces the probability of a successful simultaneous pregnancy and, if pregnancy succeeds, sibling competition for maternal milk ensues. Competition is usually won by the older sibling leading in many cases to the death of the newborn. The rate of energy transfer to pups is high in food-rich years and low in scarce ones. Because pups are more likely to be weaned as yearlings when juvenile growth rate is high than when it is...

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