Sex-biased survival predicts adult sex ratio variation in wild birds

Adult sex ratio (ASR) is a central concept in population demography and breeding system evolution, and has implications for population viability and biodiversity conservation. ASR exhibits immense interspecific variation in wild populations, although the causes of this variation have remained elusive. Using phylogenetic analyses of 187 avian species from 59 families, we show that neither hatching sex ratios nor fledging sex ratios correlate with ASR. However, sex-biased adult mortality is a significant predictor of ASR, and this relationship is robust to 100 alternative phylogenetic hypotheses, and potential ecological and life-history confounds. A significant component of adult mortality bias is sexual selection acting on males, whereas increased reproductive output predicts higher mortality in females. These results provide the most comprehensive insights into ASR variation to date, and suggest that ASR is an outcome of selective processes operating differentially on adult males and females. Therefore, revealing the causes of ASR variation in wild populations is essential for understanding breeding systems and population dynamics.     

Population regulation in the Wood Mouse Apodemus sylvaticus (L.)

Watts 1969 described the population cycle of the Wood Mouse Apodemus sylvaticus as a decline in numbers in spring, a stable period in summer and an increase phase in autumn. Aggression by adult males was considered an important aspect of population regulation limiting male survival in spring and juvenile recruitment in summer. However, recent studies of range behaviour and experimental investigations suggest that male A. sylvaticus are not relevant in population regulation and that the principal regulating factors act on female reproductive success. Female reproductive activity is determined by spatial interactions which, in turn, are determined primarily by the density of females and the proportion of female immigrants in the population. The only impact adult males may have on population size is through infanticide and competition with the females for food.     

Mating opportunity and the evolution of sex-specific mortality rates in a butterfly

Life history theory predicts that organisms should only invest resources into intrinsic components of life span to the degree that it pays off in terms of reproductive success. Here, we investigate if the temporal distribution of mating opportunities may have influenced the evolution of intrinsic mortality rates in the butterfly Pararge aegeria (Satyrinae). In this species, females mate only once and the frequency of male mating opportunities depends on the temporal emergence pattern of virgin females. As expected, in a population from Madeira where females emerge continuously throughout the year, there was no sex difference in adult life span, while in a Swedish population with synchronised female emergence, males had significantly shorter life spans compared to females. A logistic mortality model provided the best fit to the observed change in age-specific mortality and all categories reached an asymptotic mortality rate of a similar magnitude. However, the Swedish males reached this mortality plateau more rapidly than the other categories. External mortality, due to water and food limitation, affected the pattern of sex-specific mortality but males from Sweden still had higher rates of mortality compared to all other categories. We argue that selection on male longevity is likely to be weaker in Sweden because under synchronised emergence, all females emerge and mate within a short period of time, after which male reproductive value will quickly approach zero. On Madeira, however, male reproductive value decrease more slowly with age since the probability of finding a receptive female is constant over the year. Received: 29 July 1999 / Accepted: 23 August 1999     

Food limitation and demography of a migratory antelope,the white-eared kob

Summary Although it is commonly presumed that many populations of large-herbivores are limited by food availability, supporting evidence is scarce. This hypothesis was investigated in a population of over 800,000 white-eared kob in the Boma National Park region of the southern Sudan. Food availability, body condition, and mortality rates of adults and calves were measured during the dry seasons of 1982 and 1983. Sampled age distributions from both the live population and carcasses were used to calculate age-specific rates of mortality. In 1982, food supplies during the dry season were augmented by substantial rainfall, which produced regrowth of grass in areas that ordinarily had little green forage. As a result, fat reserves declined little, and rates of adult mortality showed no increasing trend. Total adult mortality was 5%. In 1983, there was no rainfall during the dry season and food intake was insufficient to meet the estimated energy requirements of kob. As a result, fat reserves declined and adult mortality rates increased fourfold. Total adult mortality was 10% (equivalent to the recruitment rate of yearling into the population). Calf mortality during the dry season was similar in both years (50%), based on field estimates of mortality rates and calf/female ratios. Lactation throughout the dry season possibly provided a buffer for calves against variations in food availability. The age structure of the live population in 1983 suggests that a drought in 1980 reduced kob numbers by 40%. These results suggest that adult survival is influenced strongly by the availability of food during the dry season. However, the duration of the dry season also plays an important role. During the dry season, declining fat reserves make an increasing proportion of the population vulnerable to mortality. As a result, even moderate droughts may lead to substantial changes in population numbers.     

Growth and body size in populations of mink frogs Rana septentrionalis from two latitudes

Age, longevity, and growth in mink frogs Rana septentrionalis from two latitudes in Quebec, Canada, were assessed by skeletochronological and back-calculation methods in order to document proximate causes for intraspecific variations in adult body size. In both study sites, females grew faster and were on average 11% larger than males. Mean age and maximal longevity were significantly higher in females than in males only in the southern populations. There was thus an interpopulation difference in the relative contribution of age and growth to sexual size dimorphisms. Sex-ratio also favored females (males sulfering higher mortality) only in the southern populations. Specimens from the northern population had higher mean ages (but not higher longevities) and were 17% larger than specimens from the southern populations. Annual growth rate appeared similar at the two study sites despite a shorter growing season at the northern locality. Maturity was reached by both sexes after 1 yr of post-metamorphic life (PML) in the southern populations but after 2 yr of PML in the northern population. There are indications that tadpoles metamorphose at larger body size at the northern locality after a prolonged larval period. It is concluded that growth rate, delayed maturity, greater mean ages, and eventually size at transformation, all contribute to the larger size of adult mink frogs at northern localities.     

Density‐independent predation affects migrants and residents equally in a declining partially migratory elk population

Migration is expected to benefit individuals through exposure to higher quality forage and reducing predation rates more than non‐migratory conspecifics. Previous studies of partially migratory ungulates (with migrant and resident individuals) have focused on bottom–up factors regulating resident and migrant segments, yet differential predation between strategies could also be a density‐dependent regulatory mechanism. Our study tested for density‐dependence in mortality, as well as mechanisms of ­bottom–up or top–down regulation in the resident and migrant portions of the partially migratory Ya Ha Tinda elk population. We tested for density dependence in adult female and juvenile survival rates, and then discriminated between predator‐ and food‐regulation hypotheses by testing for density‐dependence amongst mortality causes for adult female elk. Notably, the population declined almost 70% from near previously published estimates of carrying capacity over 10 years, providing ideal conditions to test for density dependence. In contrast to predictions, we found only weak support for density dependence in adult survival and juvenile survival. We also found few differences between migrant and resident elk in adult or juvenile survival, though juvenile survival differences were biologically significant. Predation by humans and grizzly bears was density dependent, but similar between migratory strategies. Predation by wolves was the leading known cause of mortality, yet remained constant with declining elk density equally for both migrant and resident elk, indicating wolf predation was density‐independent. Instead of being strongly regulated by food or predation, we found adult female survival was driven by density‐independent predation and climatic factors. The few differences between migratory strategies suggest equivalent fitness payoffs for migrants and residents. This population is being limited by density‐independent predation leading to declines of both migratory strategies. Our results challenge classical predator–prey theory, and call for better integration between predator–prey and migration theory.     

Long-term dynamics of fecal corticosterone in male great gerbils (Rhombomys opimus Licht.): effects of environment and social demography

We examined the relationship among seasonal characteristics of climate, food, and population demography (social structure) and fecal corticosterone (CORT) concentrations over 6 yr in adult males of an arid-adapted species, the great gerbil (Rhombomys opimus Licht., Gerbillidae, Rodentia), as a measure of chronic stress in high, low, and recovering population densities. Results showed yearly differences in the seasonal means of CORT, with the highest concentrations in the year of the highest population density. Analysis of year-specific relationships revealed a positive correlation between mean CORT and total precipitation in January and February and a negative correlation with precipitation in March. In the beginning of spring, when gerbils were in maximum reproductive effort, CORT correlated positively with the saturation of burrow systems and with the number of adult females with an adult male. A linear stepwise regression of CORT in individual males in spring seasons of all 6 yr combined after removal of year effects revealed that CORT depended positively on the number of females associated with a single male but negatively on the abundance of annual herbs. Disappearance of adult males was not related to CORT in most cases. We found no correlation between overall mortality from season to season and mean CORT in either spring (March-May) or fall. In fact, we found a highly negative correlation between mean CORT and the proportion of disappeared males at the beginning of spring. Only at the high population density when cases of probable catastrophic mortality of all adults in the group were excluded was CORT of individual males related positively to their disappearance during the summer drought. Our results suggest that desert rodents with irregular population fluctuations are more sensitive to suppression by external factors than by density-dependent mortality mediated by stress. The favorable feeding and climatic conditions may have compensated for density-dependent increases of CORT and the negative effects it might have had on survival.     

Sex-biased juvenile survival in a bird with extreme size dimorphism, the great bustard Otis tarda

We explored sex-biased mortality patterns in a species showing the most extreme sexual dimorphism among birds, the great bustard Otis tarda . Between 1991 and 2005 we studied juvenile and immature survival in a sample of 361 great bustards radio-tagged at two different populations in Spain, Villafáfila and Madrid. Mortality decreased with age, from high rates during the first year (0.70), to 0.10 in the second year. Using the known-fate model in program MARK we found that monthly survival increased throughout the first year. Offspring showing higher body mass at marking, i.e. those hatched earlier in the season and those with better body condition, survived in higher proportion. This was probably related to the earlier breeding dates of more experienced mothers, as well as to the observed decrease in food availability as the season progresses. Monthly survival estimates were higher in females than in males, which suggests that juvenile males are more vulnerable to reduced food availability and other factors due to their much faster growth rates. The proportion of non-natural deaths increased with age, and was higher in the Madrid population, where illegal hunting and collision with powerlines showed a high incidence. The male-biased mortality found in young birds in this study explains the female-biased population sex ratios observed in great bustard populations. The different degrees of incidence of human-induced causes of mortality found between both populations studied suggest that such differences may contribute to the variation observed in the adult sex ratio among populations.     

Stochastic density effects on adult fish survival and implications for population fluctuations

The degree to which population fluctuations arise from variable adult survival relative to variable recruitment has been debated widely for marine organisms. Disentangling these effects remains challenging because data generally are not sufficient to evaluate if and how adult survival rates are regulated by stochasticity and/or population density. Using unique time series for a largely unexploited reef fish, we found both population density and stochastic food supply impacted adult survival. The estimated effect of variable survival on adult abundance (both mean and variability) rivalled that of variable recruitment. Moreover, we show density‐dependent adult survival can dampen impacts of stochastic recruitment. Thus, food variability may alter population fluctuations by simultaneously regulating recruitment and compensatory adult survival. These results provide an additional mechanism for why intensified density‐independent mortality (via harvest or other means) amplifies population fluctuations and emphasises need for research evaluating the causes and consequences of variability in adult survival.     

Fetal origins of hyperphagia, obesity, and hypertension and postnatal amplification by hypercaloric nutrition

Environmental factors and diet are generally believed to be accelerators of obesity and hypertension, but they are not the underlying cause. Our animal model of obesity and hypertension is based on the observation that impaired fetal growth has long-term clinical consequences that are induced by fetal programming. Using fetal undernutrition throughout pregnancy, we investigated whether the effects of fetal programming on adult obesity and hypertension are mediated by changes in insulin and leptin action and whether increased appetite may be a behavioral trigger of adult disease. Virgin Wistar rats were time mated and randomly assigned to receive food either ad libitum (AD group) or at 30% of ad libitum intake, or undernutrition (UN group). Offspring from UN mothers were significantly smaller at birth than AD offspring. At weaning, offspring were assigned to one of two diets [a control diet or a hypercaloric (30% fat) diet]. Food intake in offspring from UN mothers was significantly elevated at an early postnatal age. It increased further with advancing age and was amplified by hypercaloric nutrition. UN offspring also showed elevated systolic blood pressure and markedly increased fasting plasma insulin and leptin concentrations. This study is the first to demonstrate that profound adult hyperphagia is a consequence of fetal programming and a key contributing factor in adult pathophysiology. We hypothesize that hyperinsulinism and hyperleptinemia play a key role in the etiology of hyperphagia, obesity, and hypertension as a consequence of altered fetal development.     

Essential amino acid supplements increase muscle weight, bone mass and bone strength in adult osteoporotic rats

Protein undernutrition is known to play an important role in the pathogenesis of osteoporotic fracture in elderly. The mechanisms underlying the bone loss in protein undernutrition appeared to be related to an uncoupling between increased bone resorption and bone formation. This was associated with decreased plasma IGF-I levels, with anoestrus and decreased muscle mass. Reversibility of protein undernutrition-induced bone loss was investigated in ovariectomized adult rats, which were fed isocaloric 2.5 % casein diet (OVX2.5) for 16 weeks. Then, the animals were given a supplement of essential amino-acids in similar proportion to that of casein at doses of 2.5% (EAA2.5) or 5% (EAA5) of total food intake for an additional 16 weeks. Essential amino acid supplements increased bone mineral mass and strength in ovariectomized protein-deprived rats. EAA supplements were associated with stimulated bone formation and reduced bone resorption, with increment of plasma IGF-I and of limb muscle mass weight. These results suggest that nutritional intervention with essential amino acid supplements can increase bone mineral mass, bone strength and muscle mass in osteoporotic rats possibly by correcting IGFI status.     

Growth and mortality in juvenile European wild rabbits (Oryctolagus cuniclus)

Factors influencing the growth and mortality patterns of young were examined in a small population of European wild rabbits. Juveniles were trapped and tagged for individual identification soon after their emergence from underground nests. Age was estimated from weight on first capture and survivorship determined from the last date individuals were observed. Differences in growth rates of young born throughout the season and between the three years of the study were related to population density. Maternal social status had no significant effect on post-emergence growth rates of kittens, or on offspring survival. Young born in females' first litters of the season, in March, had significantly lower mortality than subsequent litters. Cats and stoats were regularly observed on the study site and are likely to have been major agents of juvenile mortality. There was no difference in the growth rates of the sexes, and correspondingly non-breeding adult males and females were of similar body weight. From their first days above ground juvenile males suffered significantly higher mortality than females. Differential mortality may be linked to previously reported sex differences in behaviour if the greater activity of young males made them more vulnerable to predation or attack by neighbouring territorial conspecifics.     

Population dynamics of the whitefly,Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), in Java,Indonesia, with special reference to spatio-temporal changes in the quantity of food resources

Six monitoring plots were established in the northern part of West Java, Indonesia, to clarify the factors that influence population fluctuations of the whitefly,Bemisia tabaci. Yellow sticky traps were used to monitor adult populations. To examine the relationship between the population fluctuations and the quantity of food resources, the quantity of food resources ofB. tabaci (i.e. soybean and mung bean) was also investigated routinely in each subdistrict where the experimental plot was located. The maximum number of adults per trap in each sowing season showed a similar noticeable seasonal trend in all experimental plots. Climatic factors did not have a major role to play in population fluctuations. The number of adults per trap in each experimental plot tended to increase when the quantity of food resources in each subdistrict increased. The changes of the quantity of food resources seemed to influence the population fluctuations. It is suggested that the operation of regulatory processes in population density is influenced largely by both the distance between habitat patches and the amplitude of temporal fluctuations of the quantity of food resources. An explanation for why the time lag between fluctuations of the quantity of food resources and the number of individuals ofB. tabaci occurred is discussed. Based on their flight ability and intrinsic rate of natural increase,B. tabaci is considered not to be a serious pest in the environment where the host plants are grown discontinuously in time and space.     

Distribution and population density of the bushcricket Decticus verrucivorus in a damp-meadow biotope

Summary A population of Decticus verrucivorus (Orthoptera: Tettigoniidae) was studied in the field over two successive years. The majority of insects accumulate only in certain parts of a diverse habitat. In this area the mean population density of adult Decticus verrucivorus was 3.1/100 m². Analysis of nearest neighbour distances shows that males have a clumped distribution in the undisturbed habitat. It is concluded that larval distribution reflects the distribution of oviposition sites but that the distribution of adult males, though also depending on the distribution of natural resources and the climatic conditions, is produced mainly by the acoustic interactions with other males. The distribution of adult females does not appear to be determined by the acoustic behaviour of the males. The role of environmental factors such as the availability of food, humidity and temperature, and of the acoustic communication within this species, is discussed with respect to the problem of the high site fidelity of Decticus verrucivorus.     

Environmental factors and life histories of isolated river stocks of brown trout (Salmo trutta m. fario) in Søre Osa river system,Norway

Synopsis The brown trout in the Søre Osa river system are isolated from the lake above by a dam. They are generally of a small size, but their condition coefficient is high. They reach sexual maturity at an early age and have a short life span. The substratum in the Søre Osa mainly consists of large stones covered with a dense mossy vegetation that creates good fish cover. A high biomass of zoobenthos gives an adequate food supply throughout the year. The trout spawn mainly in tributaries, where the water level is unpredictable, some years being too low to permit successful spawning for the migrating individuals in the stock. The life history of these fish depends on the amount of food available in the habitat and on reproduction in unstable environments where density-independent mortality factors appear important. Many young migrate to the main stream during their first year. In the tributaries, there is an excess of resident males and it is hypothesized that a population structure with small resident males and large migratory males is maintained by partial inbreeding and kin selection. In the main stream the trout are larger and they have higher growth rates in the upper part than farther downstream. This is probably so because the food particles that are flushed down from the lake above provide a better supply of zoobenthic food.     

Spatial patterns and density dependence in the dynamics of a roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) population in central Italy

The demography of roe deer living in a mountain area of central Italy was studied from 1995 to 1999 with 104 radio-tagged animals, including fawns and adults of both sexes. From spring surveys we estimated population density by mark-resighting (average: 53.8dž.8 individuals km^-2) and found an average fawn/doe ratio of 0.75ǂ.4. The fawn/doe ratio was negatively correlated to density suggesting density-dependent regulation in this population. Using culled and net-trapped individuals we evaluated the dressed body weights of adult males (23.1ǃ.0 kg) and females (22.0ǃ.0 kg), which indicated a low level of sexual dimorphism. The potential litter size (1.44ǂ.1 embryos) depended on female body weight and a threshold of 20.9ǃ.4 kg separated adult females carrying one or two embryos. Both fawn (0.38ǂ.07) and adult survival (0.90ǂ.07) were evaluated from radiotagged individuals and no gender effect was observed in either age class. During the study period we recorded a population decline in one part of the study area and an increasing fawn mortality, which was attributed to the spreading of an enteropathogenic desease. The study revealed an unexpected spatial structure in population dynamics at a scale of few square kilometres. In the two studied subareas, which are very close and ecologically similar, we documented significant differences in several demographic parameters: females in the subarea with the highest deer density produced smaller litters and allocated their reproductive effort preferentially to males, which is consistent with the hypothesis that local resource competition determines sex allocation in roe deer. The importance of spatial variability with respect to roe deer demography was overlooked in previous studies and our results raise new interesting research questions relative to the study of population equilibria which are also relevant for the management of this important game species.     

Factors affecting recruitment age and recruitment probability in the Western Gull Larus occidentalis

We studied recruitment age (i.e. the age that a gull bred for the first time) and the probability that an adult reproduced during its life in three cohorts of Western Gulls Larus occidentalis breeding on southeast Farallon Island. We examined the effects of sex, hatching date, hatching order, brood size and annual variation in food supply during the recruitment period. Modal recruitment ages of males and females were 4 and 5 years, respectively, a difference that was significant. None of the factors examined, including hatching date, hatching order and brood size, had a significant effect on recruitment age. The probability of breeding in males who experienced low food supply in their fourth and fifth years was lower than that of males who experienced favourable food supply in their fourth and fifth years. Recruitment probability (the proportion of the sample group that recruited among those available to recruit during a given year) of 4- and 5-year-old males was lower in the food-poor 1983 El Nino than in years of favourable food supply. The only variable affecting recruitment probability of females was age: recruitment probability was greater among 5-year-olds than 4-year-olds. Recruitment age in males was more sensitive to annual variation in food supply, probably because females have more difficulty recruiting due to an excess of adult females in the population and because of the different roles in reproduction in which males are primarily responsible for provisioning chicks and maintaining the territory.     

Trade-offs of predation and foraging explain sexual segregation in African buffalo

1. Many studies have investigated why males and females segregate spatially in sexually dimorphic species. These studies have focused primarily on temperate zone ungulates in areas lacking intact predator communities, and few have directly assessed predation rates in different social environments. 2. Data on the movement, social affiliation, mortality and foraging of radio-collared African buffalo (Syncerus caffer) were collected from 2001-06 in the Kruger National Park, South Africa. 3. The vast majority of mortality events were due to lion (Panthera leo) predation, and the mortality hazard associated with being an adult male buffalo in a male-only 'bachelor' group was almost four times higher than for adult females in mixed herds. The mortality rates of adult males and females within mixed herds were not statistically different. Mortality sites of male and female buffalo were in areas of low visibility similar to those used by bachelor groups, while mixed herds tended to use more open habitats. 4. Males in bachelor groups ate similar or higher quality food (as indexed by percentage faecal nitrogen), and moved almost a third less distance per day compared with mixed herds. As a result, males in bachelor groups gained more body condition than did males in breeding herds. 5. Recent comparative analyses suggest the activity-budget hypothesis as a common underlying cause of social segregation. However, our intensive study, in an area with an intact predator community showed that male and female buffalo segregated by habitat and supported the predation-risk hypothesis. Male African buffalo appear to trade increased predation risk for additional energy gains in bachelor groups, which presumably leads to increased reproductive success.     

Is the future female for turtles? Climate change and wetland configuration predict sex ratios of a freshwater species

Climate change and land-use change are leading drivers of biodiversity decline, affecting demographic parameters that are important for population persistence. For example, scientists have speculated for decades that climate change may skew adult sex ratios in taxa that express temperature-dependent sex determination (TSD), but limited evidence exists that this phenomenon is occurring in natural settings. For species that are vulnerable to anthropogenic land-use practices, differential mortality among sexes may also skew sex ratios. We sampled the spotted turtle (Clemmys guttata), a freshwater species with TSD, across a large portion of its geographic range (Florida to Maine), to assess the environmental factors influencing adult sex ratios. We present evidence that suggests recent climate change has potentially skewed the adult sex ratio of spotted turtles, with samples following a pattern of increased proportions of females concomitant with warming trends, but only within the warmer areas sampled. At intermediate temperatures, there was no relationship with climate, while in the cooler areas we found the opposite pattern, with samples becoming more male biased with increasing temperatures. These patterns might be explained in part by variation in relative adaptive capacity via phenotypic plasticity in nest site selection. Our findings also suggest that spotted turtles have a context-dependent and multi-scale relationship with land use. We observed a negative relationship between male proportion and the amount of crop cover (within 300 m) when wetlands were less spatially aggregated. However, when wetlands were aggregated, sex ratios remained consistent. This pattern may reflect sex-specific patterns in movement that render males more vulnerable to mortality from agricultural machinery and other threats. Our findings highlight the complexity of species' responses to both climate change and land use, and emphasize the role that landscape structure can play in shaping wildlife population demographics.     

Investigations on the feeding habits of the rocky-shore miteHyadesia fusca (Acari: Astigmata: Hyadesiidae): diet range, food preference, food quality, and the implications for distribution patterns

Within the food web of estuarine and marine rocky shore ecosystems phytophagous mites of terrestrial and marine origin constitute an important part as grazers on algae and as a food source for certain arthropods, especially zoophagous mites. This investigation deals with the feeding biology ofHyadesia fusca taking as an example a population located on an artificial rocky shore of the middle Weser estuary in Northern Germany. The species is characterized by a broad diet range; in feeding experiments diatoms, lichens, detritus as well as blue, red and green algae were accepted. Even analyses of faecal pellets produced by field specimen suggest a non-specific feeding habit. However, the influence of certain diets on mortality, offspring number and rearing success showed that the food quality differs significantly. The most suitable food, the UlvaceaeBlidingia, was clearly preferred in a series of pairwise choice tests. These findings correlate with the vertical zonation of the field population i.e.: higher population densities in the vegetation zone dominated byBlidingia. It can be concluded that in addition to abiotic factors food supply could play an important role for distribution patterns of phytophagous mites.