Cornelia Kraus

Physiological suppression of sexual function of subordinate males: a subtle form of intrasexual competition among male sifakas (Propithecus verreauxi)?

In contrast to most anthropoid primates, sifakas (Propithecus verreauxi), like many group-living lemurs, exhibit a number of features that deviate from predictions of sexual selection theory. Despite a promiscuous mating system, they lack sexual dimorphism, suggesting that physical combat plays only a minor role in intrasexual competition for receptive females. In this study, we investigated the hypothesis that socioendocrinological mechanisms contribute to suppression of reproductive function of subordinate males. For that purpose, 10 male sifakas from five social groups were observed for 669 focal animal hours for 4 months, including the mating season, in Western Madagascar. Concomitantly 315 fecal samples of these animals were collected and the concentration of immunoreactive testosterone was quantified hy enzymeimmunoassay procedures. Clear dominance relationships existed among coresident males. Testosterone levels of dominant males were significantly higher than those of subordinates during, as well as outside, the mating season. Additionally, the increase in testosterone levels prior to the mating season was more pronounced for dominant than for subordinate males. These findings are in accordance with the hypothesis of suppression of sexual function of subordinate males, probably providing dominant males with ani advantage in sperm competition. If reproductive success is mainly determined by this nonagonistic form of intrasexual competition, the results of this study contribute an important piece to the puzzle of lacking sexual dimorphism in P. verreauxi.

Influence of reproductive season and rank on fecal glucocorticoid levels in free-ranging male Verreaux's sifakas (Propithecus verreauxi)

Studies in anthropoid primates and other mammals suggest that reproductive season, rank, reproductive skew, aggression received, and social support are the major factors influencing glucocorticoid output. In which way these are also affecting adrenal function in lemurid primates has been studied rarely. Here, we examine the influence of reproductive season and rank on glucocorticoid output in male sifakas (Propithecus verreauxi), a species characterized by high breeding seasonality, a hierarchy among males and extreme reproductive skew towards dominant males. We established a fecal assay for non-invasively monitoring adrenal activity and collected 315 fecal samples during the reproductive and birth season from 10 male sifakas living in 5 groups in Western Madagascar. We found a significant effect of season on glucocorticoid output, with males exhibiting higher fecal glucocorticoid levels during the reproductive compared to the birth season in conjunction with an increase in overall aggression rates during the former period. Moreover, our data indicate a significant effect of rank on adrenocortical activity with dominant males exhibiting higher glucocorticoid levels than subordinate males in the reproductive season. However, dominant males did not differ significantly in rates of initiated or received aggression and rates of affiliative behavior from subordinates but showed significantly lower rates of submission. Given their highly formalized dominance relationships, we conclude that higher glucocorticoid output in dominant males during the 4-month reproductive season is likely related to higher energetic demands necessary to cope with the challenges of male reproduction rather than to physical demands of increased fighting frequency to maintain dominance status. High rank in sifakas may thus carry high costs, which, however, may be outweighed by monopolization of almost all paternities. In sum, our data generally support the findings on the relationship between environmental and social factors and glucocorticoid output found in non-lemurid primates.

The Size–Life Span Trade-Off Decomposed: Why Large Dogs Die Young

Large body size is one of the best predictors of long life span across species of mammals. In marked contrast, there is considerable evidence that, within species, larger individuals are actually shorter lived. This apparent cost of larger size is especially evident in the domestic dog, where artificial selection has led to breeds that vary in body size by almost two orders of magnitude and in average life expectancy by a factor of two. Survival costs of large size might be paid at different stages of the life cycle: a higher early mortality, an early onset of senescence, an elevated baseline mortality, or an increased rate of aging. After fitting different mortality hazard models to death data from 74 breeds of dogs, we describe the relationship between size and several mortality components. We did not find a clear correlation between body size and the onset of senescence. The baseline hazard is slightly higher in large dogs, but the driving force behind the trade-off between size and life span is apparently a strong positive relationship between size and aging rate. We conclude that large dogs die young mainly because they age quickly.

Large males dominate: ecology, social organization, and mating system of wild cavies, the ancestors of the guinea pig

Ecological factors differently affect male and female animals and thereby importantly influence their life history and reproductive strategies. Caviomorph rodents are found in a wide range of habitats in South America and different social and mating systems have evolved in closely related species. This permits to study the impact of ecological factors on social evolution. In this study, we investigated the social organization and the mating system of the wild cavy (Cavia aperea), the ancestor of the domestic guinea pig, in its natural habitat in Uruguay. Based on our laboratory investigations, we expected a polygynous system with large males controlling access to females. Results from radiotelemetry and direct observations showed that females occupied small stable home ranges which were largely overlapped by that of one large male, resulting in a social organization of small harems. In some cases, small satellite males were associated with harems and intermediate-sized roaming males were occasionally observed on the study site. However, microsatellite analyses revealed that offspring were exclusively sired by large males of the same or neighboring harems, with a moderate degree of multiple paternity (13–27%). Thus, the mating system of C. aperea can be described as polygynous and contrasts with the promiscuous organization described for other species of cavies (Cavia magna, Galea musteloides and Microcavia australis) living under different ecological conditions. Our findings stress the strong impact of environmental factors on social evolution in Caviomorphs as resource distribution determines female space use and, thereby, the ability of males to monopolize females.

The costs of risky male behaviour: sex differences in seasonal survival in a small sexually monomorphic primate

Male excess mortality is widespread among mammals and frequently interpreted as a cost of sexually selected traits that enhance male reproductive success. Sex differences in the propensity to engage in risky behaviours are often invoked to explain the sex gap in survival. Here, we aim to isolate and quantify the survival consequences of two potentially risky male behavioural strategies in a small sexually monomorphic primate, the grey mouse lemur Microcebus murinus: (i) most females hibernate during a large part of the austral winter, whereas most males remain active and (ii) during the brief annual mating season males roam widely in search of receptive females. Using a 10-year capture–mark–recapture dataset from a population of M. murinus in Kirindy Forest, western Madagascar, we statistically modelled sex-specific seasonal survival probabilities. Surprisingly, we did not find any evidence for direct survival benefits of hibernation—winter survival did not differ between males and females. By contrast, during the breeding season males survived less well than females (sex gap: 16%). Consistent with the ‘risky male behaviour’ hypothesis, the period for lowered male survival was restricted to the short mating season. Thus, sex differences in survival in a promiscuous mammal can be substantial even in the absence of sexual dimorphism.

Senescence or selective disappearance? Age trajectories of body mass in wild and captive populations of a small-bodied primate

Classic theories of ageing consider extrinsic mortality (EM) a major factor in shaping longevity and ageing, yet most studies of functional ageing focus on species with low EM. This bias may cause overestimation of the influence of senescent declines in performance over condition-dependent mortality on demographic processes across taxa. To simultaneously investigate the roles of functional senescence (FS) and intrinsic, extrinsic and condition-dependent mortality in a species with a high predation risk in nature, we compared age trajectories of body mass (BM) in wild and captive grey mouse lemurs (Microcebus murinus) using longitudinal data (853 individuals followed through adulthood). We found evidence of non-random mortality in both settings. In captivity, the oldest animals showed senescence in their ability to regain lost BM, whereas no evidence of FS was found in the wild. Overall, captive animals lived longer, but a reversed sex bias in lifespan was observed between wild and captive populations. We suggest that even moderately condition-dependent EM may lead to negligible FS in the wild. While high EM may act to reduce the average lifespan, this evolutionary process may be counteracted by the increased fitness of the long-lived, high-quality individuals.

Reproduction and growth in a precocial small mammal, Cavia magna

Abstract Small mammals usually produce large litters of altricial young, resulting in high reproductive rates. In contrast, cavies give birth to few precocial young after a long gestation. The price of this reproductive strategy is a low intrinsic rate of natural increase. We investigated if the patterns of reproduction in a wild population of Cavia magna are consistent with the hypotheses that cavies can increase their reproductive output by breeding aseasonally and by maturing extremely early. We collected data on reproduction and growth by capture–recapture during a 26-month field study in a wetland in Uruguay, and from a laboratory population founded with individuals from the same region. Among the Caviinae, C. magna is particularly precocial, with individual neonates weighing on average 18% of maternal mass. Reproduction was mostly seasonal, with the main birth season starting at the end of September (austral spring) and extending until May in 1999 and February in 2000, respectively, with only a few females reproducing during the 1st but not the 2nd austral winter. Individual females produced on average 3 litters per year. Some females born in early spring conceived successfully between the age of 30 and 45 days, similar to females in the laboratory. The remainder of the 1st spring cohort and females of subsequent birth cohorts delayed reproduction until the following spring. Body condition and growth rates were highest in the spring, declined through the year, and varied between years, and may be the proximate factors determining whether an adult female or a juvenile initiates breeding. Breeding opportunistically whenever conditions allow might partly compensate for the low reproductive rate of cavies.

Levels and mechanisms of behavioural variability

The behaviour of animals is the result of adaptations and constraints. To what extent a particular behaviour pattern can be attributed to the relative strengths of these two forces is crucial with respect to understanding the evolution of behaviour. Moreover, in the still young history of the study of animal behaviour, different conceptual approaches have placed very different emphases on the residual behavioural variability beyond adaptations and constraints. In this chapter, we retrace some of these paradigm shifts, offer an overview of different hierarchical levels at which behavioural variability occurs and summarise some of the mechanisms that generate or constrain it. Above the species level, phylogenetic constraints often limit behavioural variability because of a functional relationship between taxonwide life history traits and behaviour, but the exact nature of their underlying mechanisms remains obscure. Phylogenetic constraints exist at different taxonomic levels, and, as several examples from studies of primate behaviour indicate, they are also common in animals with relatively advanced cognitive abilities in which social learning is common. We therefore emphasise the importance of acknowledging the existence of such constraints in behavioural analyses. We also decompose behavioural variability further into variation within species, among individuals and within individuals over time and highlight some of the mechanisms responsible for generating and maintaining this variability. Our review suggests that the specific evolutionary history of a taxon will set the stage at which levels variability can arise, and that cognitive abilities appear to create surprisingly little additional freedom for behavioural variability.

The stress of growing old: sex- and season-specific effects of age on allostatic load in wild grey mouse lemurs

Chronic stress [i.e. long-term elevation of glucocorticoid (GC) levels] and aging have similar, negative effects on the functioning of an organism. Aged individuals’ declining ability to regulate GC levels may therefore impair their ability to cope with stress, as found in humans. The coping of aged animals with long-term natural stressors is virtually unstudied, even though the ability to respond appropriately to stressors is likely integral to the reproduction and survival of wild animals. To assess the effect of age on coping with naturally fluctuating energetic demands, we measured stress hormone output via GC metabolites in faecal samples (fGCM) of wild grey mouse lemurs (Microcebus murinus) in different ecological seasons. Aged individuals were expected to exhibit elevated fGCM levels under energetically demanding conditions. In line with this prediction, we found a positive age effect in the dry season, when food and water availability are low and mating takes place, suggesting impaired coping of aged wild animals. The age effect was significantly stronger in females, the longer-lived sex. Body mass of males but not females correlated positively with fGCM in the dry season. Age or body mass did not influence fGCM significantly in the rainy season. The sex- and season-specific predictors of fGCM may reflect the differential investment of males and females into reproduction and longevity. A review of prior research indicates contradictory aging patterns in GC regulation across and even within species. The context of sampling may influence the likelihood of detecting senescent declines in GC functioning.

Mama’s boy: sex differences in juvenile survival in a highly dimorphic large mammal, the Galapagos sea lion

In many mammals, early survival differs between the sexes, with males proving the more fragile sex [“Fragile male (FM) hypothesis”], especially in sexually dimorphic species where males are the larger sex. Male-biased allocation (MBA) by females may offset this difference. Here, we evaluate support for the FM and MBA hypotheses using a dataset on Galapagos sea lions (Zalophus wollebaeki). We statistically model sex-specific survival as it depends on body mass and environmental conditions (sea surface temperature, SST, a correlate of marine productivity) at three developmental stages, the perinatal phase (1st month), the main lactation period (1st year), and the weaning period (2nd year). Supporting the FM hypothesis, we found that, early in life (1st month), at equal birth mass, males survived less well than females. During the remainder of the first year of life, male survival was actually less sensitive to harsh environmental conditions than that of females, contradicting the FM hypothesis and supporting the MBA hypothesis. During the second year of life, only male survival suffered with high SSTs as predicted by the FM hypothesis. At each developmental stage, observed survival rates were almost equal for both sexes, suggesting that mothers buffer against the inherent fragility of male offspring through increased allocation, thereby masking the differences in survival prospects between the sexes.