Intraspecific allometry of neonatal size in the antarctic fur seal (Arctocephalus galapagoensis)

Summary Intraspecific allometry of pup mass as a function of maternal mass was analysed in Antarctic fur seals (Arctocephalus gazella), a highly dimorphic species. The allometric exponent was 0.6, much lower than in interspecific comparisons. Slopes were the same for mother-son pairs as for mother-daughter pairs, but adjusted means were significantly higher for the former, indicating higher reproductive effort of mothers of sons. The correlation of maternal mass with pup mass explained 59% of the variance in female pup mas but only 18% of that in male pup mass. Females appeared to produce sons which were as big as possible. Relative pup mass decreased with increasing maternal mass indicating a lower reproductive effort of bigger and presumably older mothers, contrary to expectations from life history theory. Sex ratio of pups showed no relation to maternal mass.     

Maternal investment and sex-allocation in the Galapagos fur seal,Arctocephalus galapagoensis

Summary Maternal investment and sex-allocation were measured in a large, sexually dimorphic mammal, the Galapagos fur seal (Arctocephalus galapagoensis). The sex ratio at birth was 1.06. Males were always heavier than females and, at least initially, grew faster. Growth was variable from year to year suggesting energetic constraints on maternal investment. Sucking time conrrelated with milk intake. Mothers suckled yearling and 2-year-old sons more than daughters of the same age. Age at weaning appeared to be the same in both sexes or even slightly greater in males. No sex differences was found in mortality prior to weaning or in post-weaning dispersal. Birth rates of females with yearlings or 2-year-olds were significantly lower than those of females with no dependent young. Mothers invested more in sons than in daughters until weaning. It is unlikely that higher post-weaning investment in daughters balances the higher pre-weaning investment in sons. Data on sex ratio at birth, different growth rates, and weaning age of the sexes are typical of otariid seals as a group. The results of this study fit Maynard Smith's (1980) model of the evolution of sex allocation better than Fisher's (1930).     

Weaning in the guinea pig (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): Who decides and by what measure?

Offspring should be selected to influence maternal effort to maximize their own fitness, whereas mothers are selected to limit investment in present progeny. In mammals, this leads to a conflict over the amount of milk provided and the timing of weaning. The intensity and time course of such conflict has so far mostly been investigated experimentally in altricial rodents. However, it is expected that offspring options for conflict will depend on developmental state. We therefore investigated in the highly precocial domestic guinea pig (Cavia aperea f. porcellus) who decides over nursing performance and weaning and how pup state influences these decisions. Specifically, we tested whether a threshold mass of pups predicts weaning time. By exchanging older litters against neonates and vice versa, we produced a situation in which females differed in lactational stage from the cross-fostered pups. Our results indicate that females decide about the timing of weaning, as cross-fostered younger pups were weaned at a much younger age than controls and older pups benefited from continuing lactation of foster mothers. Growth rates did not differ in the treatment groups, and different weaning ages resulted in differing weaning mass refuting the hypothesis that weaning is based on a threshold mass of offspring. This constitutes clear evidence that in a precocial rodent, the guinea pig, decisions about maternal care are primarily determined by maternal state and little influenced by pup state despite the extreme precociality of offspring. We suggest that precocial pups show little resistance to early weaning when food is abundant, as they reach sufficient nutritional independence by the middle of lactation to enable independent survival.     

Sibling competition and cooperation in mammals: challenges,developments and prospects

Many vertebrates grow up in the company of same or different-age siblings, and relations among them can be expected to significantly influence individual life histories and the development of individual morphological, physiological, and behavioral phenotypes. Although studies in birds still dominate and have stimulated most theoretical considerations, the increasing number of mammalian studies promises to broaden our understanding of this complex field by enabling interesting comparisons with the rather different bird system. It therefore seems timely to bring together recent studies of sibling relations in mammals and to demonstrate what these can offer in the way of fresh insights. In this brief review, intended to accompany a series of papers on a diverse range of mammals, we outline the current state of sibling research in mammals, comparing it to the better studied birds. Most obviously, in mammals, mother and young are in much closer contact during early life than in birds, and siblings can influence each other’s development as well as the mother’s physiology while still in utero. During nursing, mammalian young also encounter a very different feeding situation to bird siblings. These contrasts should help stimulate further debate, as well as provide further opportunities to study the relative importance of maternal versus sibling effects on individual development. Finally, we discuss the need to balance studies of sibling competition and conflict with a consideration of the benefits accruing to individuals from sibling presence and the need for long-term studies of the influence of early sibling relations on individual development and life histories. This contribution represents the introduction to the special issue “Sibling competition and cooperation in mammals”.     

Lactation patterns of pinnipeds are not explained by optimization of maternal energy delivery rates

Within pinnipeds, phocids and otariids show differing maternal care strategies. Phocids rear young out of body stores in a yearly cycle with a single stay ashore when the mother fasts while lactating, whereas otariids provision their young by repeated foraging trips to sea alternating with brief stays ashore where they suckle their young. In a previous optimality model, these differences have been interpreted as adaptations based on differing energy requirements of large (phocid) and smaller (otariid) species, and the time budget of the large elephant and the much smaller Antarctic fur seal were correctly predicted. Our refined model—extended to pinniped species of all sizes—predicts lactation strategies to shift from attendance cycles to 1-year cycles with increasing body mass and provides an explanation for the finding that phocid pups are weaned at lower relative mass than otariid pups. However, other predictions do not correspond to empirical findings. In particular, the model does not explain the behavior of large otariids and small phocids. Thus, maternal metabolic requirements alone appear insufficient to explain observed lactation patterns. In the light of our results, we discuss more generally the scope and limitations of optimality models when applied in a comparative framework to a group of related species.     

Site fidelity of male Galápagos sea lions: a lifetime perspective

Knowledge about the distribution of resources can lead to the development of spatial preferences and long-term site fidelity. Individuals are expected to choose sites that best suit their needs. However, dominant individuals restrict movements of less competitive ones. Accordingly, one may expect spatial preferences to differ with regard to individual characteristics and to change over time. We investigated lifetime changes of site fidelity patterns with regard to reproductive success in male Galápagos sea lions (Zalophus wollebaeki). Showing a high degree of natal philopatry in the first 2 years of life, non-territorial males pass through a stage of fidelity to their natal colony where they develop preferences for areas outside prime breeding areas. Variation in the degree of spatial preferences was not associated with age or size, characteristics linked to an individual’s dominance status. For non-territorial males, roaming proved to be an adequate strategy to gain reproductive success. Only the most competitive males established territories in areas preferentially visited by females. They had a high probability to return to breeding areas where they successfully reproduced in previous seasons. Overall, the results reveal lifetime changes in site fidelity with regard to male status. The degree of site fidelity observed within the colony suggests familiarity and thus a high degree of tolerance among individuals using the same areas. This seems to facilitate attendance in the colony and thus the possibility to prospect for oestrus females.     

Foraging strategies of the marine iguana,Amblyrhynchus cristatus

Summary Two foraging strategies were found in marine iguanas (Amblyrhynchus cristatus); (1) subtidal feeding: the animals swam out to sea and dived for algae in the subtidal zone; (2) intertidal feeding: the animals foraged around low tide in the intertidal zone on more or less exposed algae. Most marine iguanas were very consistent in their foraging strategy and so could be classified as subtidal feeders (SFs) or intertidal feeders (IFs). Feeding strategy was weight-related (Fig. 1), not sexspecific. Animals 1,200 g were IFs, animals >1,800 g SFs. Some iguanas in between followed a mixed foraging strategy. SFs foraged independently of the tides, IFs always around low tide (Figs. 2, 3). Feeding time patterns of IFs and SFs are described (Table 1). Sea motion seemed to have little effect on the foraging pattern of SFs, but strongly influenced that of IFs (Fig. 2). The smaller a marine iguana, the faster it cooled when immersed in water (Fig. 4). The difference between water temperature and core temperature of animals returning from foraging was significantly greater in IFs than SFs (Fig. 5). SFs swimming in very cold water regulated their body temperature to prevent excessive cooling. Possible costs and benefits of the two foraging strategies are discussed. Only part of a marine iguana population lives really amphibiously and only ca. 5% of a 24 h day is spent close to or in the water. All social activities, including mating, take place on land. These life history characteristics preclude those adaptations to an amphibious way of life that would at the same time reduce the iguanas' ability to be maximally active at their typical terrestrial body temperature of 35° C.     

Female aggression and male peace-keeping in a cichlid fish harem: conflict between and within the sexes in Lamprologus ocellatus

Summary Conflicts of interest within and between the sexes are important processes leading to variability in mating systems. The behavioral interactions mediating conflict are little documented. We studied pairs and harems of the snail-shell inhabiting cichlid fish Lamprologus ocellatus in the laboratory. Due to their larger size, males controlled the resource that limited breeding: snail shells. Males were able to choose among females ready to spawn. Females were only accepted if they produced a clutch within a few days of settling. When several females attempted to settle simultaneously the larger female settled first. Females were least aggressive when guarding eggs. Secondary females were more likely to settle when the primary female was guarding eggs. In established harems females continued to be aggressive against each other. The male intervened in about 80% of female aggressive interactions. Male intervention activity correlated with the frequency of aggression among the females in his harem. The male usually attacked the aggressor and chased her back to her own snail shell. When a male was removed from his harem, aggression between females increased immediately and usually the secondary female was expelled by the primary female within a few days. Time to harem break-up was shorter the more mobile the primary females' young were and did not correlate with the size difference between harem females. Male L. ocellatus interfere actively in female conflict and keep the harem together against female interests. Female conflict presumably relates to the cost of sharing male parental investment and to the potential of predation by another female's large juveniles on a female's own small juveniles.Correspondence to: F. Trillmich     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).