Temperature-dependent sex-biased embryo mortality in a bird

Sex ratios have important evolutionary consequences and are often biased by environmental factors. The effect of developmental temperature on offspring sex ratios has been widely documented across a diverse range of taxa but has rarely been investigated in birds and mammals. However, recent field observations and artificial incubation experiments have demonstrated that the hatching sex ratio of a megapode, the Australian brush-turkey (Alectura lathami), varied with incubation temperature; more females hatched at high incubation temperatures and more males hatched at low temperatures. Here, we investigated the causes of this temperature-dependent sex-biasing system. Molecular sexing of chicks and embryos confirmed that male embryo mortality was greater at high temperatures while female embryo mortality is greater at low temperatures, with mortality in both sexes similar at intermediate incubation temperatures. Temperature-dependent sex-biased embryo mortality represents a novel mechanism of altering sex ratios in birds. This novel mechanism, coupled with the unique breeding biology of the brush-turkey, offers a potentially unparalleled opportunity in which to investigate sex allocation theory in birds.     

Reproductive ageing and sexual selection on male body size in a wild population of antler flies (Protopiophila litigata)

Little is known about the importance of trade-offs between ageing and other life history traits, or the effects of ageing on sexual selection, particularly in wild populations suffering high extrinsic mortality rates. Life history theory suggests that trade-offs between reproduction and somatic maintenance may constrain individuals with higher initial reproductive rates to deteriorate more rapidly, resulting in reduced sexual selection strength. However, this trade-off may be masked by increased condition dependence of reproductive effort in older individuals. We tested for this trade-off in males in a wild population of antler flies (Protopiophila litigata). High mating rate was associated with reduced longevity, as a result of increased short-term mortality risk or accelerated ageing in traits affecting viability. In contrast, large body size was associated with accelerated ageing in traits affecting mating success, resulting in reduced sexual selection for large body size. Thus, ageing can affect sexual selection and evolution in wild populations.     

Diversification rates and the latitudinal gradient of diversity in mammals

The latitudinal gradient of species richness has frequently been attributed to higher diversification rates of tropical groups. In order to test this hypothesis for mammals, we used a set of 232 genera taken from a mammalian supertree and, additionally, we reconstructed dated Bayesian phylogenetic trees of 100 genera. For each genus, diversification rate was estimated taking incomplete species sampling into account and latitude was assigned considering the heterogeneity in species distribution ranges. For both datasets, we found that the average diversification rate was similar among all latitudinal bands. Furthermore, when we used phylogenetically independent contrasts, we did not find any significant correlation between latitude and diversification parameters, including different estimates of speciation and extinction rates. Thus, other factors, such as the dynamics of dispersal through time, may be required to explain the latitudinal gradient of diversity in mammals.     

Division of labour influences the rate of ageing in weaver ant workers

The evolutionary theory of ageing predicts that the timing of senescence has been primarily shaped by the extrinsic mortality rate, which causes selection intensity to decline over time. One difficulty in testing the evolutionary theory of ageing is that extrinsic mortality risk is often confounded with body size and fecundity, which may also directly affect lifespan. Social insects with a pronounced division of labour between worker castes provide a unique opportunity to study the direct effect of extrinsic mortality on the evolution of ageing rates independently of body size, reproductive effort and genetic configuration. In the weaver ant, Oecophylla smaragdina, the major (large) workers perform the risky tasks outside the nest, while the minor (small) workers stay within the highly protected arboreal nest. Hence, this pronounced division of labour is associated with high differences in extrinsic mortality risks. The evolutionary theory of ageing predicts that the minor workers should have a longer intrinsic lifespan than the major workers. In line with this prediction, we found that in a protected environment the minor workers lived significantly longer than the major workers did. Hence, the ageing rate appears to have been moulded by variation in the extrinsic mortality rate independently of size, reproductive effort and genetic configuration.     

Predicting natural mortality rates of plants and animals

Understanding the factors that control the mortality rates of species in their natural environments is important for understanding the structure and dynamics of populations, communities and ecosystems. Here, we test a model of natural mortality that yields explicit, quantitative predictions based on the constraints of body size and temperature on individual metabolism. Extensive field data from plants, invertebrates, fish, birds and mammals indicate that much of the heterogeneity in rates of natural mortality can be predicted, despite the many extrinsic sources of mortality in natural systems. These results suggest that common 'rule(s)' govern mortality rates in ecological communities for organisms as diverse as plants and animals.     

Life-history variation within a taxon: a comparative analysis of birds and mammals

Most studies on life-history evolution discuss the necessity of distinguishing between extrinsic and intrinsic sources of variability in life-history traits. I use log/log plots of yearly neonate production in daugters (b) versus adult mortality (Ma) for 75 bird species and 88 mammal species to compare graphically life-history "fields" arranged by these selective forces along a "slow-fast continuum". Under the assumptions of steady-state and linear relationship between adult mortality and reproductive effort, as well as between juvenile survival and relative neonate weight, it is possible to place additional axes in the two-dimentional plot, and to predict covariations among demographic and individual growth traits. The functional regression analysis shows, that the assumptions are completely fulfilled, at least for birds, but mammals show nonlinear relationship between adult mortality and reproductive effort. This can be explained by peculiarities of metabolism and parental care in small mammals with high reproductive output. Hence, for birds the axis of relative neonate weight approximately coincides in direction with the juvenile survivorship axis, but this is not a case for mammals. In both taxa, the relative neonate weight is an invariant in relation to fecundity and adult mortality (but not in relation to adult body weight). This important feature, together with other intrinsic (energetic and phylogenetic) constraints, explains well-documented close covariations among traits, even when the effect of body size is factored out. It is argued that life-history and body size variations in birds and mammals mainly depend on a pattern of temporal resource deficiency, although this impact cannot be separated from that of extrinsic juvenile mortality.     

Lack of age-related mosaic loss of W chromosome in long-lived birds

Females and males often exhibit different survival in nature, and it has been hypothesized that sex chromosomes may play a role in driving differential survival rates. For instance, the Y chromosome in mammals and the W chromosome in birds are often degenerated, with reduced numbers of genes, and loss of the Y chromosome in old men is associated with shorter life expectancy. However, mosaic loss of sex chromosomes has not been investigated in any non-human species. Here, we tested whether mosaic loss of the W chromosome (LOW) occurs with ageing in wild birds as a natural consequence of cellular senescence. Using loci-specific PCR and a target sequencing approach we estimated LOW in both young and adult individuals of two long-lived bird species and showed that the copy number of W chromosomes remains constant across age groups. Our results suggest that LOW is not a consequence of cellular ageing in birds. We concluded that the inheritance of the W chromosome in birds, unlike the Y chromosome in mammals, is more stable.     

Life history patterns in birds and mammals and their evolutionary interpretation

Summary It is argued that allometric principles account for most of the observed variation in the life history patterns amongst birds. To test this contention it is shown that traits such as incubation time, growth rates, age at first reproduction, lifespan, clutch weight and egg weight all scale to body weight with exponents similar to those found for analogous traits in mammals. It is then shown that most of the variation amongst bird taxa and between birds and mammals based on body weight allometry can be explained by variations in brain size, body temperature and metabolic rate, consistent with theories of growth and ageing derived from mammalian studies. Finally, it is suggested that the evidence for life histostory allometry is sufficiently strong that it argues for a more epigenetic view of life history patterns and their evolution than is generally conceded in most adaptation theories.     

Is group size related to longevity in mammals?

Life-history theory predicts that reduced extrinsic risk of mortality should increase species longevity over evolutionary time. Increasing group size should reduce an individual''s risk of predation, and consequently reduce its extrinsic risk of mortality. Therefore, we should expect a relationship between group size and maximum longevity across species, while controlling for well-known correlates of longevity. We tested this hypothesis using a dataset of 253 mammal species and phylogenetic comparative methods. We found that group size was a poor predictor of maximum longevity across all mammals, as well as within primates and rodents. We found a weak but significant group-size effect on artiodactyl longevity, but in a negative direction. Body mass was consistently the best predictor of maximum longevity, which may be owing to lower predation risk and/or lower basal metabolic rates for large species. Artiodactyls living in large groups may exhibit higher rates of extrinsic mortality because of being more conspicuous to predators in open habitats, resulting in shorter lifespans.     

Embryo development and ageing in birds and mammals

The rate of ageing is a genetically influenced feature of an individual's life history that responds to selection on lifespan. Various costs presumably constrain the evolution of prolonged life, but these have not been well characterized and their general nature is unclear. The analyses presented here demonstrate a correlation among birds and mammals between rates of embryonic growth and ageing-related mortality, which are quantified by the exponents of fitted power functions. This relationship suggests that rapid early development leads to accelerated ageing, presumably by influencing some aspect of the quality of the adult individual. Although the mechanisms linking embryo growth rate and ageing are not known, a simple model of life-history optimization shows that the benefits of longer life can be balanced by connected costs of extended development.     

Evolution of longevity through optimal resource allocation

Models of life history evolution predict optimal traits of a simplified organism under various environmental conditions, but they at most acknowledge the existence of ageing. On the other hand, genetic models of ageing do not consider the effects of ageing on life histroy traits other than fecundity and longevity. This paper reports the results of a dynamic programming model which optimizes resource allocation to growth, reproduction and somatic repair. A low extrinsic (environmentally caused) mortality rate and high repair efficiency promote allocation to repair, especially early in life, resulting in delayed ageing and low growth rates, delayed maturity, large body size and dramatic enhancement of survival and maximum lifespan. The results are generally consistent with field, comprative and experimental data. They also suggest that the relationships between maximum lifespan and age at maturity and body size observed in nature may be by-products of optimal allocation strategies.     

Extrinsic Mortality Effects on Reproductive Strategies in a Caribbean Community

Extrinsic mortality is a key influence on organisms’ life history strategies, especially on age at maturity. This historical longitudinal study of 125 women in rural Domenica examines effects of extrinsic mortality on human age at maturity and pace of reproduction. Extrinsic mortality is indicated by local population infant mortality rates during infancy and at maturity between the years 1925 and 2000. Extrinsic mortality shows effects on age at first birth and pace of reproduction among these women. Parish death records show huge historical variation in age-specific mortality rates. The infant mortality rate (IMR) in the early 1920s was low, increased dramatically beginning in 1929, and reached a maximum in the 1950s, at which point IMR declined steadily to its present low rate. The mortality rate early in life showed a quadratic association with age at first birth. Women who experienced conditions of low IMR early in life reproduced relatively late in life. Those born into moderately high levels of infant mortality tended to reproduce earlier than those born at low levels. At very high infant mortality levels early in life, women went on to delay reproduction until relatively late, possibly as a result of somatic depletion and energetic stress associated with the conditions that lead to high IMR. Population mortality rates at age of maturity also showed a quadratic association with age at first birth. The pace of reproduction, estimated as number of surviving offspring controlled for maternal age, showed a similar quadratic effect. There were complex interactions between population mortality rates in infancy and at maturity. When extrinsic mortality was high during infancy, extrinsic mortality later in life had little effect on timing of first birth. When extrinsic mortality was low to moderate in infancy, extrinsic mortality later in life had significant effects on adult reproduction. I speculate that these effects are mediated through development of personality facets associated with reproduction.     

Population dynamics of large and small mammals

We offer an evaluation of the Caughley and Krebs hypothesis that small mammals are more likely than large mammals to possess intrinsic population regulating mechanisms. Based on the assumption that intrinsic regulation will be manifest via direct density-dependent feedbacks, and extrinsic regulation via delayed density-dependent feedbacks, we fit autoregressive models to 30 time series of abundance for large and small mammals to characterize their dynamics. Delayed feedbacks characterizing extrinsic mechanisms, such as trophic-level interactions, were detected in most time series, including both small and large mammals. Spectral analyses indicated that the effect of such delayed feedbacks on the variability in population growth rates differed with body size, with large mammals exhibiting predominantly reddened and whitened spectra in contrast with predominantly blue spectra for small mammals. Large mammals showed less variance and more stable dynamics than small mammals, consistent with, among other factors, differences in their potential population growth rates. Patterns of population dynamics in small versus large mammals contradicted those predicted by the Caughley and Krebs hypothesis.     

Predation by Bears Drives Senescence in Natural Populations of Salmon

Classic evolutionary theory predicts that populations experiencing higher rates of environmentally caused (“extrinsic”) mortality should senesce more rapidly, but this theory usually neglects plausible relationships between an individual''s senescent condition and its susceptibility to extrinsic mortality. We tested for the evolutionary importance of this condition dependence by comparing senescence rates among natural populations of sockeye salmon (Oncorhynchus nerka) subject to varying degrees of predation by brown bears (Ursus arctos). We related senescence rates in six populations to (1) the overall rate of extrinsic mortality, and (2) the degree of condition dependence in this mortality. Senescence rates were determined by modeling the mortality of individually-tagged breeding salmon at each site. The overall rate of extrinsic mortality was estimated as the long-term average of the annual percentage of salmon killed by bears. The degree of condition dependence was estimated as the extent to which bears killed salmon that exhibited varying degrees of senescence. We found that the degree of condition dependence in extrinsic mortality was very important in driving senescence: populations where bears selectively killed fish showing advanced senescence were those that senesced least rapidly. The overall rate of extrinsic mortality also contributed to among-population variation in senescence-but to a lesser extent. Condition-dependent susceptibility to extrinsic mortality should be incorporated more often into theoretical models and should be explicitly tested in natural populations.     

Annual Survival and Turnover Rates of an Afrotropical Robin in a Fragmented Forest

Whether or not subdivided populations persist in fragmented landscapes primarily depends on how well individuals can survive within discrete habitat patches. Using data from six capture–recapture sessions, survival probabilities of the white-starred robin were estimated in seven indigenous forest patches in the highly fragmented Taita Hills forests, SE Kenya. We found no significant differences in survival probability either among fragment-size categories (large 135 ha, medium 95 ha, and small 2–8 ha) or between adult and first-year birds. However, males had a higher probability of survival from one year to the next than females. Turnover rates of adult birds were higher for females than males, but also higher in the medium and small patches than in the large one within each sex. That survival probability was similar among fragments, but turnover rates differed denoted that different processes caused extirpation from the patches. We suggest that mortality associated with dispersal was probably a more important cause of extirpation than within-patch mortality in the largest habitat, which had the lowest turnover rates. Conversely, high within-patch mortality, for instance due to predation during incubation, could have been more important in the smaller, more disturbed habitats. These results lend support to the proposition that avian conservation efforts should be focussed both at the landscape level to improve connectivity between fragments and reduce mortality during dispersal, and at the patch level to exclude other mortality sources such as nest predation.     

Why fly? The possible benefits for lower mortality

In adult endotherms—birds and mammals—mortality rates are lower in flying than non-flying species. Rates vary with size, but allowing for this they are higher in penguins than flying birds, and in rodents than bats. These observations are most simply explained in terms of predation since flight, allowing movement in three dimensions, increases the chances of escape.
There are various problems in obtaining and comparing data on rates of mortality and this partly explains a wide scattering of points. Nevertheless the main results are statistically significant, mostly at P<0.001. Amongst birds, there are several other significant differences. Mortality rates are lower at lower latitudes, and in aquatic compared to terrestrial species, with cliff-nesters having lower rates than other aquatic birds and co-operative breeders than other terrestrial ones. No latitudinal effect was detected in mammals. The exceptionally low rates of mortality in bats and swifts are attributed to their being particularly hard to catch.     

The maximum oxygen consumption and aerobic scope of birds and mammals: getting to the heart of the matter

Resting or basal metabolic rates, compared across a wide range of organisms, scale with respect to body mass as approximately the 0.75 power. This relationship has recently been linked to the fractal geometry of the appropriate transport system or, in the case of birds and mammals, the blood vascular system. However, the structural features of the blood vascular system should more closely reflect maximal aerobic metabolic rates rather than submaximal function. Thus, the maximal aerobic metabolic rates of birds and mammals should also scale as approximately the 0.75 power. A review of the literature on maximal oxygen consumption and factorial aerobic scope (maximum oxygen consumption divided by basal metabolic rate) suggests that body mass influences the capacity of the cardiovascular system to raise metabolic rates above those at rest. The results show that the maximum sustainable metabolic rates of both birds and mammals are similar and scale as approximately the 0.88 +/- 0.02 power of body mass (and aerobic scope as approximately the 0.15 +/- 0.05 power), when the measurements are standardized with respect to the differences in relative heart mass and haemoglobin concentration between species. The maximum heart beat frequency of birds and mammals is predicted to scale as the -0.12 +/- 0.02 power of body mass, while that at rest should scale as -0.27 +/- 0.04.     

Insights from comparative analyses of aging in birds and mammals

Many laboratory models used in aging research are inappropriate for understanding senescence in mammals, including humans, because of fundamental differences in life history, maintenance in artificial environments, and selection for early aging and high reproductive rate. Comparative studies of senescence in birds and mammals reveal a broad range in rates of aging among a variety of taxa with similar physiology and patterns of development. These comparisons suggest that senescence is a shared property of all vertebrates with determinate growth, that the rate of senescence has been modified by evolution in response to the potential life span allowed by extrinsic mortality factors, and that most variation among species in the rate of senescence is independent of commonly ascribed causes of aging, such as oxidative damage. Individuals of potentially long‐lived species, particularly birds, appear to maintain high condition to near the end of life. Because most individuals in natural populations of such species die of aging‐related causes, these populations likely harbor little genetic variation for mechanisms that could extend life further, or these mechanisms are very costly. This, and the apparent evolutionary conservatism in the rate of increase in mortality with age, suggests that variation in the rate of senescence reflects fundamental changes in organism structure, likely associated with the rate of development, rather than physiological or biochemical processes influenced by a few genes. Understanding these evolved differences between long‐lived and short‐lived organisms would seem to be an essential foundation for designing therapeutic interventions with respect to human aging and longevity.     

Seasonal variation in the metabolism-temperature relation of House Sparrows (Passer domesticus) in KwaZulu-Natal,South Africa

Many birds exhibit considerable phenotypic flexibility in metabolism to maintain thermoregulation or to conserve energy. This flexibility usually includes seasonal variation in metabolic rate. Seasonal changes in physiology and behavior of birds are considered to be a part of their adaptive strategy for survival and reproductive success. House Sparrows (Passer domesticus) are small passerines from Europe that have been successfully introduced to many parts of the world, and thus may be expected to exhibit high phenotypic flexibility in metabolic rate. Mass specific Resting Metabolic Rate (RMR) and Basal Metabolic Rate (BMR) were significantly higher in winter compared with summer, although there was no significant difference between body mass in summer and winter. A similar, narrow thermal neutral zone (25–28 °C) was observed in both seasons. Winter elevation of metabolic rate in House Sparrows was presumably related to metabolic or morphological adjustments to meet the extra energy demands of cold winters. Overall, House Sparrows showed seasonal metabolic acclimatization similar to other temperate wintering passerines. The improved cold tolerance was associated with a significant increase in VO₂ in winter relative to summer. In addition, some summer birds died at 5 °C, whereas winter birds did not, further showing seasonal variation in cold tolerance. The increase in BMR of 120% in winter, compared to summer, is by far the highest recorded seasonal change so far in birds.