Proximate and ultimate explanations of mammalian sex allocation in a marsupial model

Offspring sex ratios in mammals vary in potentially adaptive yet unpredictable ways. An integrative approach that simultaneously examines proximate and ultimate explanations of mammalian sex ratios would greatly advance the field. We examined the importance of maternal glucose and stress hormones for offspring sex (male or female) as mechanisms associated with the Trivers–Willard and the local resource competition hypotheses of sex allocation. We tested this framework in a marsupial mammal, the tammar wallaby (Macropus eugenii). Mothers that were better able to maintain body condition over the driest part of the year, a presumptive proxy for local resource availability, were more likely to produce daughters (the philopatric sex), consistent with local resource competition. Maternal glucose was correlated with offspring sex, but in the opposite direction than we predicted—higher maternal glucose was associated with female pouch young. These patterns, however, were not consistent across the 2 years of our study. Maternal stress hormone metabolites measured from fecal samples did not predict glucose or offspring sex. A causative glucose mechanism may underlie an adaptive strategy for mothers with high local resources (high glucose) to produce philopatric daughters that will benefit from inheriting resource access. Examining species-specific relationships between glucose and offspring sex across mammals could provide crucial insight into the disparate ecological and selective pressures faced by mammals with respect to offspring sex ratio.     

Are faecal hormone levels linked to winter progression,diet quality and social rank in young ungulates ? An experiment with white-tailed deer (<Emphasis Type="Italic">Odocoileus virginianus</Emphasis>) fawns

Hormones play a central role in the physiology and behaviour of animals. The recent development of noninvasive techniques has increased information on physical and social states of individuals through hormone measurements. The relationships among hormones, life history traits and behaviours are, however, still poorly known. For the first time, we evaluated natural winter glucocorticoid and testosterone levels in young ungulates in relation to winter progression, diet quality and social rank. Overwinter, levels of glucocorticoid and testosterone decreased, possibly due to the decline of fawns’ body mass. The relationships between hormone levels and diet quality were surprising: Fawns fed the control diet presented higher glucocorticoid and lower testosterone levels then fawns fed the poor diet, suggesting that control fawns faced a higher nutritional stress than those on the poor diet. Similarly to other studies on social mammals, we found no relationship between faecal glucocorticoid levels and social rank, suggesting that social stress was similar for dominant and subordinate fawns during winter. Testosterone levels were not correlated to social rank as found previously in groups of individuals forming stable social hierarchies and maintaining stable dominance relationships. The simultaneous suppression of glucocorticoid and testosterone levels suggests for the first time that young ungulates present a hormonal strategy to prevent fast depletion of limited proteins and fat resources during winter.     

Maternal investment in mountain baboons and the hypothesis of reduced care

It has been argued that female mammals should terminate expensive forms of infant care earlier as habitat quality declines. More recently it has been shown that among a variety of mammalian species, early termination of care is also associated with highly favourable conditions. In this paper we present data on maternal investment decisions among baboons (Papio cynocephalus ursinus) inhabiting the Drakensberg Mountains of South Africa, and compare these with data from East African baboon studies. Mothers in the mountain habitat face a set of environmental conditions where the problem of resource allocation to offspring is expected to be particularly acute. We begin by using the model of Altmann (1980) of maternal time budgets to demonstrate that mountain baboon mothers experience greater perturbations to their activity budgets while suckling than do mothers in other populations. They also provide consistently greater levels of care to their infants and do so in the absence of any form of overt conflict over access to the nipple. Although this investment results in a relative lengthening of the interbirth interval (IBI), it is accompanied by relatively higher infant survival. We argue that factors that influence the maternal strategy adopted by mountain baboons include slow infant growth rates and a lack of predation in the habitat which influences probability of offspring survival beyond the immediate postnatal period. We suggest that both “care-dependent” sources of mortality (e.g. female reproductive condition, the amount of milk transferred to offspring) as well as “care independent” sources of mortality (e.g. predation, infectious disease) should be considered in studies of parental investment. Received: 26 May 1997 / Accepted after revision: 9 August 1997     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

The effects of size-dependent predation risk on the interaction between behavioral and life history traits in a stream-dwelling isopod

In field surveys, laboratory observations and field-based assays of behavior, I examined the effects of size-dependent predation risk on the interaction between size at reproductive maturity and maternal care behavior in the stream-dwelling isopod, Lirceus fontinalis. L. fontinalis exhibit population-specific sizes at reproductive maturity which result in population differences in predation risk during the adult phase. Females from streams containing salamander larvae (that prefer small prey) mature at large sizes and then become relatively safe from predation. Females from streams containing fish (that consume all size classes of prey equally) mature at small sizes and remain at risk. I tested whether these differences in expected survival were reflected in the behavior of females during the maternal phase (i.e., the period during which females exhibit maternal care). Female L. fontinalis carry developing juveniles inside a brood pouch. I simulated predatory attacks on gravid female L. fontinalis from the different population types and found that female behavior correlated with population differences in risk. When “attacked”, females from streams with predatory fish (that experience high risk to adult females) released juveniles from the brood pouch, whereas females from populations with predatory salamander larvae (that pose relatively little risk to adult females) did not release juveniles. I discuss the results with reference to the joint evolution of behavioral and life history traits. Received: 6 March 1996 /Accepted after revision: 12 August 1996     

Does predation risk,through moon phase and predator cues,modulate food intake,antipredatory and physiological responses in wood mice (Apodemus sylvaticus)?

Predation influences the ecology and behaviour of prey species and it is well known that the risk of predation affects prey’s decision making. We investigated whether predation risk through moon phase and exposure to the faecal odour of a natural predator, the red fox Vulpes vulpes, affect feeding behaviour and physiological response in wood mice (Apodemus sylvaticus). Antipredatory response was studied by live trapping under new and full moon in odourless control areas and areas experimentally manipulated with red fox fresh faeces. Food intake by individuals was determined as the amount of bait remaining in each trap and the physiological response was measured non-invasively analysing faecal corticosterone metabolites (FCM). Traps treated with faeces of red fox were the most avoided, and this avoidance was more significant during full moon. Food intake by wood mice varied according to the moon phase being significantly lower under full moon nights. We found sex, breeding condition and weight of individuals explaining the variation found in FCM concentrations, but no changes in FCM levels due to moon phase or exposure to red fox faeces were detected. These results indicate that wood mice avoid red fox faecal odour and this antipredatory response as well as feeding behaviour are significantly influenced by moon phase. However, no physiological response was found due to predation risk suggesting that wood mice do not take these predation cues enough reliable to experience physiological changes.     

Patterns of brood division and an absence of behavioral plasticity in a neotropical passerine

Studies of parental behavior in various habitats provide an opportunity to gain insight into how different environments may mold strategies of parental care. Brood division by parents has been hypothesized to occur facultatively within and among species. Brood division occurs when each parent cares for specific offspring within a brood. We studied brood division in a neotropical passerine, the western slaty antshrike (Thamnophilus atrinucha). Our results present a unique picture of a highly specialized example of avian brood division. Division was a fixed behavioral pattern in the population studied: all broods divided by fledging and remained divided during the entire post-fledging period. Brood division before fledging, a previously unreported phenomenon, occurred in 40% of nests observed. Parents that preferentially fed a certain offspring (defined as their focal offspring) in the nest fed the same individual after fledging. Each parent fed only its focal offspring in broods of one and two. The male parent cared for the heavier offspring and the first offspring to leave the nest. Siblings were segregated spatially during the time of highest predation risk. These observations suggest that a consistently high risk of predation on offspring has favored initial spatial segregation and inflexibility of brood division behavior in this species. Factors other than predation risk alone may explain the observed patterns of long-term, perfect brood division. Because high predation is common and relatively predictable in the tropics, selection for fixed brood division may be stronger in tropical birds than in the temperate zone.     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Prey-mediated effects of drought on condition and survival of a terrestrial snake

Drought can have severe ecological effects and global climate-change theory predicts that droughts are likely to increase in frequency and severity. Therefore, it is important that we broaden our understanding of how drought affects not only individual species, but also multitrophic interactions. Here we document vegetation and small-mammal abundance and associated patterns of Texas ratsnake (Elaphe obsoleta) body condition and survival before, during, and after a drought in central Texas, USA. Vegetation (grass and forbs) height and small-mammal capture rates were two times greater in wet years compared to the drought year. The decline of small mammals (the snakes' principal prey) during the drought was associated with a drop in ratsnake body condition, consistent with reduced food intake. During the drought, snake mortality also increased 24%. Although higher snake mortality was attributable to predation and road mortality rather than being a direct result of starvation, an increase in risk-prone behavior by foraging snakes probably increased their exposure to those other mortality factors. Drought conditions lasted only for 21 months, and vegetation, small-mammal abundance, and snake condition had returned to pre-drought levels within a year. Although estimates of snake population size were not available, it is likely that substantially more than a year was required for the population to return to its previous size.     

Sex-biased maternal expenditure in Rocky Mountain bighorn sheep

Summary We examined maternal energy expenditure in Rocky Mountain bighorn sheep to test the hypothesis (Trivers and Willard 1973) that individual sons in polygynous mammals should obtain a larger maternal subsidy. In accord with theory, males weighed more at birth (an index of relative prenatal expenditure), tended to suckle more between 40 and 100 days of age (an index of relative postnatal expenditure) and imposed greater delays on their mothers' return to estrus in the subsequent breeding season (an index of relative total preweaning expenditure). Mothers rather than offspring appeared to have primary control over postnatal expenditure. The effect of maternal reproductive effort on return to estrus was cumulative over a period of 3–5 years so that ewes making relatively large expenditures gave birth progressively later. Ewe and lamb mortality was associated with late conception in the previous rut. Thus, differential reproductive effort by ewes in this population appeared to entail a fitness cost involving reductions in both offspring and maternal survivorship. Correspondence to: J.T. Hogg at the present address     

Intergenerational effects of climate generate cohort variation in lizard reproductive performance

An evaluation of the link between climate and population dynamics requires understanding of climate effects both within and across generations. In ectothermic vertebrates, demographic responses to climate changes should crucially depend on balancing needs for heat and water. Here, we studied how temperature and rainfall regimes experienced before and during adulthood influenced reproductive performances (litter size, offspring size, and survival) in a natural population of the live-bearing common lizard, Lacerta vivipara, monitored continuously from 1989 to 2004. Rainfall regime, but not temperature, had both immediate and delayed effects on these reproductive performances. Rainfall during the first month of life was positively correlated with juvenile survival. Females experiencing more rainfall during gestation produced smaller neonates that showed greater survival when controlling for the positive effect of body size on survival. Furthermore, females that experienced heavier rainfall when in utero produced fewer but longer neonates during adulthood. These demographic effects of rainfall on adult reproductive traits may come from maternal effects of climate conditions and/or from delayed effects of rainfall on the environment experienced early in life. Irrespective of the precise mechanism, however, this study provides evidence of intergenerational climate effects in natural populations of an ectothermic vertebrate.     

Mixed sex allocation strategies in a polytocous mammal,the house mouse (<Emphasis Type="Italic">Mus musculus</Emphasis>)

The issue of adaptive adjustment of offspring sex ratio (proportion of male births) in polytocous mammals, producing several offspring per litter, is controversial because females of these species can maximize their fitness mainly by adjusting offspring number. To address this issue, we examined the effect of maternal condition at mating, experimentally decreased by pre-mating food restriction, on the sex ratio variation in 137 female mice. We tested two basic sex allocation hypotheses plausible for polytocous mammals: (1) the Myers hypothesis, predicting that cheaper sex should be favored in poor environmental conditions to maximize offspring number; and (2) the Williams hypothesis, predicting maximum fitness returns by adjusting size- and sex-specific composition of the litter according to the maternal condition. The food-restricted mothers produced larger litters with a higher proportion of cheaper daughters than the control mothers. By contrast, the control mothers optimized size and sex composition of the litter according to their weight at mating. In addition, the offspring of the food-restricted mothers suffered less from pre-weaning mortality than those of the control mothers. Therefore, when comparing the groups, the Myers hypothesis had a general significance while the Williams hypothesis was plausible only for the control mothers. Furthermore, some of the food-restricted mothers partly coped with the pre-mating food restriction and increased the proportion of sons in the litter with the increasing maternal weight loss (during the period of food restriction). The sex ratio variation was thus a result of three sex allocation strategies depending on the maternal condition at mating.     

Is the antipredatory response in behaviour reflected in stress measured in faecal corticosteroids in a small rodent?

Predation risk has been shown to alter various behaviours in prey. Risk alters activity, habitat use and foraging, and weight decrease might be a consequence of that. In mammals, studies on physiological measures affected by risk of predation, other than weight, are rare. We studied in two separate laboratory experiments foraging, hoarding behaviour and expression of stress measured non-invasively from the faeces in the bank vole (Clethrionomys glareolus), a common boreal rodent. Voles were exposed to predation risk using odours of the least weasels (Mustela nivalis nivalis). Distilled water served as control. In the first experiment, we found that foraging effort, measured as sunflower seeds taken from seed trays filled with sand, was significantly lower in trays scented with weasel odour. Both immediate consumption of seeds and hoarding were affected negatively by the weasel odour. Females hoarded significantly more than males in autumn. In the second experiment, the negative effect of weasel odour on foraging was consistent over a 3-day experiment, but the strongest effect was observed in the first night. Foraging increased over the time of the experiment, which might reflect either energetic compensation during a longer period of risk, predicted in the predation risk allocation hypothesis, or habituation to the odour-simulated risk. Despite decreased foraging under predation risk, stress measured as corticosteroid metabolite concentration in vole faeces was not affected by the weasel odour treatment. In conclusion, we were able to verify predation-risk-mediated changes in the foraging effort of bank voles but no physiological stress response was measured non-invasively, probably due to great individual variation in secretion of stress hormones.     

Molecular Genetic Consequences of a Population Bottleneck Associated with Reintroduction of the Mauna Kea Silversword (Arg yroxiphium sandwicense ssp. sandwicense [Asteraceae])

The endangered Mauna Kea silversword (Argyroxiphium sandwicense ssp. sandwicense) has experienced a severe decline in distribution and abundance because of predation by alien ungulates. By the late 1970s only a small remnant natural population persisted on the Mauna Kea volcano on the Island of Hawaii. The Hawaii Department of Land and Natural Resources, Division of Forestry and Wildlife, initiated an outplanting program in the 1970s to promote recovery of A. sandwicense ssp. sandwicense. Intermittent outplanting since 1973 has generated an outplanted population of over 450 plants on Mauna Kea, but the program has unintentionally resulted in a major population bottleneck. All plants in the outplanted population appear to be first- or subsequent-generation offspring of only two maternal founders from the natural population. Genetic variation in the natural and outplanted populations was assessed for 90 random amplified polymorphic DNA loci. Eleven loci were detectably polymorphic in the natural population, whereas only three loci were detectably polymorphic in the outplanted population. Thus, the population bottleneck has been accompanied by a 73% reduction in the level of detectable polymorphism. In addition, for eight loci, the population bottleneck has resulted in the loss of the marker allele in the outplanted population. A management strategy involving manual pollen transfer has recently been implemented to incorporate additional founders from the natural population into the outplanting program. As a supplement to the outplanting program, the strategy also includes a program promoting direct seedling establishment following manual pollen transfer. Incorporating additional founders may serve to overcome the legacy of the population bottleneck, especially if founder representation can be equalized.     

污染物对鱼类的甲状腺激素干扰效应及其作用机制

环境中可以影响生物体甲状腺激素合成、运输、作用和代谢等过程的化学污染物称为甲状腺激素干扰物(TDCs),TDCs被认为是继环境雌激素之后最重要的一类内分泌干扰物.鱼类甲状腺的结构、甲状腺激素的转运和功能等与哺乳动物有较大差别.与哺乳动物相比,污染物干扰鱼类甲状腺激素的研究还较为缺乏.在介绍鱼类甲状腺结构、功能以及甲状腺激素在鱼体内动态过程的基础上,分析了污染物对鱼类的甲状腺激素干扰效应及其作用机制,探讨了今后鱼类甲状腺激素干扰研究的主要方向.污染物能对鱼类甲状腺激素水平、相关酶活性及甲状腺结构等产生直接影响;同时,污染物还可以通过干扰鱼类甲状腺系统对由甲状腺激素调节的重要生理过程如生长、繁殖和发育等产生间接影响.污染物主要通过干扰鱼类甲状腺激素的合成与分泌、转运、清除以及与甲状腺激素受体(TR)的相互作用等机制对鱼类产生不利影响.在污染物对鱼类甲状腺激素干扰的研究中,今后应重点关注环境中"新型"卤化有机污染物、鱼类早期发育过程与甲状腺激素干扰效应的关系、TR介导机制以及TDCs的筛选方法等.     

Egg testosterone affects wattle color and trait covariation in the ring-necked pheasant

Transfer of maternal hormones to the eggs is a major source of offspring phenotypic variation. The developmental and organizational effects of egg hormones can extend into adulthood and affect behavioral and morphological traits involved in sexual and reproductive behavior, with important consequences for offspring fitness. In this study, we injected testosterone (T) in egg albumen of captive ring-necked pheasant (Phasianus colchicus) eggs. We then assessed the consequences for chick growth, cell-mediated immunity, and multiple male secondary sexual traits at maturity by comparison with a control group. We also compared the covariation between traits in the two experimental groups. We found that control males had redder wattles than males from T-injected eggs, suggesting that attractiveness and reproductive success of the offspring might vary depending on maternal transfer of T to the eggs. T treatment also modified the covariation between cell-mediated immunity and wattle coloration and between the area of the wattle and the expression of another secondary sexual trait, the ear tufts. These effects are likely to translate into fitness differences among the offspring if mate acquisition depends on the simultaneous expression of several traits that are differentially affected by the same maternal contribution. Maternal effects mediated by egg hormones might affect the fitness of the offspring not only by directional modification of phenotypic traits, but also by facilitating or inhibiting their covariation. This suggests the possibility that female choice based on the relative expression of multiple secondary sexual traits exerts a pressure on how maternal transfer of androgens contributes to developmental programs.     

Transgenerational plasticity in the sea: context-dependent maternal effects across the life history

Maternal effects can have dramatic influences on the phenotype of offspring. Maternal effects can act as a conduit by which the maternal environment negatively affects offspring fitness, but they can also buffer offspring from environmental change by altering the phenotype of offspring according to local environmental conditions and as such, are a form of transgenerational plasticity. The benefits of maternal effects can be highly context dependent, increasing performance in one life-history stage but reducing it in another. While maternal effects are increasingly well understood in terrestrial systems, studies in the marine environment are typically restricted to a single, early life-history stage. Here, I examine the role of maternal effects across the life history of the bryozoan Bugula neritina. I exposed maternal colonies to a common pollution stress (copper) in the laboratory and then placed them in the field for one week to brood offspring. I then examined the resistance of offspring to copper from toxicant-exposed and toxicant-na?ve mothers and found that offspring from toxicant-exposed mothers were larger, more dispersive, and more resistant to copper stress than offspring from na?ve mothers. However, maternal exposure history had pervasive, negative effects on the post-metamorphic performance (particularly survival) of offspring: offspring from toxicant-exposed mothers had poorer performance after six weeks in the field, especially when facing high levels of intraspecific competition. Maternal experience can have complex effects on offspring phenotype, enhancing performance in one life-history stage while decreasing performance in another. The context-dependent costs and benefits associated with maternally derived pollution resistance may account for why such resistance is induced rather than continually expressed: mothers must balance the benefits of producing pollution-resistant larvae with the costs of producing poorer performing adults (in the absence of pollution).     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.