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.
Female intra-sexual competition plays an important role in the settlement process during pair or harem formation and in established harems of Lamprologus ocellatus , a small snail shell inhabiting cichlid from Lake Tanganyika. Larger females settle first and this could partly be due to male preference for larger females as shown in simultaneous choice tests but is also due to dominance of the larger female. Smaller females were unable to settle close to a larger one. Even when snail shells were not limiting the smaller was either unable to settle or had to settle at a considerable distance. This effect was independent of prior residence. Intense female-female aggression suggests that close settlement is disadvantageous to females. Genetic analyses of maternity using microsatellite length polymorphism at five loci showed a reproductive skew between females in a harem. Additionally, it proved brood mixing in aquaria as well as in the field. Brood mixing can be detrimental to female breeding success through interbrood cannibalism if size difference of juveniles amounts to 5 mm. Territoriality of juveniles, shown even between same-sized siblings, may cause indirect mortality through earlier dispersal of young. Females rejected experimentally added larger juveniles but accepted young smaller than their own fry. Acceptance of smaller juveniles could be advantageous through a dilution of predator attacks but it also appears to induce costs since females with young at the shell do not rear another brood. Large median distances of 91 cm maintained aggressively between breeding females in the field may serve to minimize the adverse effects of breeding in a harem.
We studied the ontogeny of diving behaviour in the Galápagos fur seal (Arctocephalus galapagoensis, Heller 1904). Six-month-old seals spent less than 12% of observation time at sea and were entirely dependent on maternal milk for nutrition. Maximum dive depths for this age group averaged 5.6 m, maximum durations 50 s (N = 12). Modal swim speeds averaged 0.55 m/s; maximum observed swim speeds ranged from 0.9-2 m/s (N = 5). Six-month-old seals swam distances of 2.9 km/24 hrs on average (N = 5). No diel patterns were apparent in their diving behaviour. One-year-old fur seals were the youngest age group to show substantial diving activity, reaching maximum depths of 47.5 m and durations of 2.4 min on average (N = 21). Modal swim speeds averaged 0.9 m/s, maximum speeds ranged from 1.4-2.3 m/s (N = 6). Total distances traveled averaged 15.4 km/trip to sea. Yearlings spent 23.9% of observation time at sea, the majority (> 95%) of that at night. This corresponds to the activity pattern of adult females who dive exclusively at night. Yearlings were the youngest age group to contribute to their own nutrition through independent foraging, but were still dependent on their mothers: no yearlings were successfully weaned in this study. 18-month-old seals were better divers, spending 27.6% of observation time at sea, mostly during the night. Maximum dive depths for this age group averaged 61.1 m; durations 3.1 min (N = 11). Modal swim speeds averaged 1.4 m/s (N = 4), and were comparable to modal speeds of two adult females of 1.2 and 1.6 m/s. Maximum speeds for these juveniles ranged from 2.2-2.7 m/s (N = 4); the two adult females reached 2.8 and 4 m/s respectively. Total travel distances averaged 42 km/trip to sea (N = 4 juveniles). These distances corresponded to those covered by two adult females, 43 and 45 km/trip. Two-year-old fur seals included the youngest that were successfully weaned in this study. They spent 35% of observation time at sea, primarily at night. Maximum dive depths averaged 69.7 m, durations 3.4 min (N = 20). Adult females were the best divers in this study, spending 49.5% of observation time at sea, almost exclusively at night. Maximum dive depths averaged 106.5 m, durations 4.5 min (N = 32). During ontogeny, age was the best predictor of time spent at sea. For animals older than six months, body mass was the best predictor for maximum dive depth as well as maximum and median durations. The physiological maturation process precludes weaning before the age of one year. After that, the diving capacity of young fur seals takes a central role in the weaning process, in conjunction with the temporal patterns of relative prey accessibility.
We describe the development and characterisation of six new dinucleotide motif microsatellite loci for populations of marine iguanas (Amblyrhynchus cristatus), endemic to the Galápagos archipelago. Primers were based on microsatellite-bearing sequences and initially developed using universally labelled primers. When analysed across 5 populations (representing 150 individuals), new loci displayed, on average, high levels of genetic diversity (range: 2-13 alleles, mean: 5.73) and values of heterozygosity (range: 0.0-0.906, mean: 0.605). No consistent deviations from Hardy-Weinberg equilibrium or significant linkage disequilibrium were observed, and all loci were shown to be free of common microsatellite errors. Utilising the 13 previously available microsatellite loci for this species, we describe here four multiplex combinations for the successful amplification of 19 microsatellite loci for marine iguanas. This powerful set of highly polymorphic markers will allow researchers to explore future questions regarding the ecology, evolution, and conservation of this unique species.