Evidence suggesting that reindeer mothers allonurse according to the direct reciprocity and generalized reciprocity decision rules.

Allonursing is the nursing of the offspring of other mothers. Cooperation is an emergent property of evolved decision rules. Cooperation can be explained by at least three evolved decision rules: 1) direct reciprocity, i.e. help someone who previously helped you, 2) kin discrimination, i.e. preferentially direct help to kin than to non-kin, and 3) generalized reciprocity, i.e. help anyone if helped by someone. We assessed if semi-domesticated reindeer, Rangifer tarandus, mothers allonursed according to the decision rules of direct reciprocity, generalized reciprocity and kin discrimination over 2 years. To assess if reindeer mothers allonursed according to the direct reciprocity decision rule, we predicted that mothers should give more help to those who previously helped them more often. To assess if reindeer mothers allonursed according to the kin discrimination decision rule, we predicted that help given should increase as pairwise genetic relatedness increased. To assess if reindeer mothers allonursed according to the generalized reciprocity decision rule, we predicted that the overall number of help given by reindeer mothers should increase as the overall number of help received by reindeer mothers increased. The number of help given i) increased as the number of help received from the same partner increased in the 2012 group but not in both 2013 groups, ii) was not influenced by relatedness, and iii) was not influenced by an interaction between the number of help received from the same partner and relatedness. iv) The overall number of help given increased as the overall number of help received increased. The results did not support the prediction that reindeer mothers allonursed according to the kin discrimination decision rule. The results suggest that reindeer mothers may allonurse according to the direct reciprocity and generalized reciprocity decision rules.

Mismatch between calf paternity and observed copulations between male and female reindeer: Multiple mating in a polygynous ungulate?

In polygynous systems, such as that exhibited by reindeer Rangifer tarandus, mate choice can be difficult to disentangle from male intrasexual competition because male behavior may constrain female choice. Multiple mating may provide an avenue for female mate choice, though it is difficult to identify using behavioral estimators alone. Molecular techniques address this issue by affording ecologists an opportunity to reassess mating systems from a genetic perspective. We assessed the frequency and possible explanations for multiple mating in reindeer using a genetic approach to determine the success of observed copulations in a semi-domesticated herd in Kaamanen, Finland. Behavioral and genetic data were synthesized with population characteristics over a 7-year period to test the hypothesis that, if present, polyandry in reindeer is driven by sexual harassment from sub-dominant males. We observed multiple mating in 42% of females, with as many as 60% exhibiting multiple mating in certain years. We found no evidence that multiple mating resulted from sexual harassment by sub-dominant males, suggesting that it is likely a deliberate strategy among females. Conversion rate of copulations into paternities varied with male size, with smaller males more likely to experience mismatch than larger males. Female preference for larger males persisted despite the occurrence of multiple mating, possibly suggesting a mechanism for cryptic post-copulatory selection. We suggest further research to delineate the possible influence of cryptic post-copulatory selection and multiple mating to defend against infertility in exhausted males.

Zoo soundscape: Daily variation of low-to-high-frequency sounds.

Most studies assessing the impact of noises on zoo animal welfare did not measure sound frequencies outside of the human-hearing range (infrasounds and ultrasounds). Many nonhuman mammals can hear these frequencies, and because loud and variable soundscapes are potentially detrimental for animal welfare, this overlooked aspect of their acoustic environment could have important consequences. This study evaluated the soundscape of an urban zoo in a large frequency range (17.5-90,510 Hz) by measuring its average sound levels (Leq ) and variability (the difference between highest and lowest peaks). Sound data were collected for 24 hr in 25 locations (e.g., indoor, outdoor, near the amusement park). The soundscape was not considered problematic for animal welfare when looking at the average sound levels in most locations (<77-dB sound pressure level [SPL]), except for a few indoor areas and near the water park. Ultrasounds were rare, had low average sound levels, and were less variable in time. Infrasounds were always present and were the loudest and most variable sound frequencies. The soundscape was louder and more variable during the day and when visitors were present, suggesting that human-related activities were the sources of these augmentations. Indoor environments were generally louder than outdoor environments and touristic features; however, the water park was near 85-dB SPL during the day. On the basis of results, we suggest a series of mitigation actions to minimize noise-related stress in captive animals.

Response of reindeer mating time to climatic variability.

BACKGROUND: The breeding time of many species has changed over the past 2-3 decades in response to climate change. Yet it is a key reproductive trait that affects individual's parturition time and reproductive success, and thereby population dynamics. In order to predict how climate change will affect species' viability, it is crucial to understand how species base their reproductive efforts on environmental cues. RESULTS: By using long-term datasets of mating behaviours and copulation dates recorded since 1996 on a semi-domesticated reindeer population, we showed that mating time occurred earlier in response to weather conditions at different key periods in their annual breeding cycle. The mating time occurred earlier following a reducing snow cover in early spring, colder minimum temperatures in the last 2 weeks of July and less precipitation in August-September. CONCLUSIONS: The mediated effect of a reduced snow cover in early spring on improving individuals' pre-rut body weight through a better availability of late winter food and reduced costs of locomotion on snow would explain that mating time has occurred earlier overtime. A lower level of insect harassment caused by colder maximum temperatures in July might have caused an advance in mating time. Less precipitation in August-September also caused the mating time to occur earlier, although the direct effects of the last two weather variables were not mediated through the pre-rut body weight of individuals. As such, the causal effects of weather conditions on seasonal timing of animals are still unclear and other mechanisms than just body weight might be involved (e.g. socio-biological factors). The plastic response of reindeer mating time to climatic variability, despite supplemental feeding occurring in late April, demonstrated that environmental factors may have a greater influence on reproductive outputs than previously thought in reindeer.

The onset in spring and the end in autumn of the thermal and vegetative growing season affect calving time and reproductive success in reindeer.

A developing trophic mismatch between the peak of energy demands by reproducing animals and the peak of forage availability has caused many species' reproductive success to decrease. The match-mismatch hypothesis (MMH) is an appealing concept that can be used to assess such fitness consequences. However, concerns have been raised on applying the MMH on capital breeders such as reindeer because the reliance on maternal capita rather than dietary income may mitigate negative effects of changing phenologies. Using a long-term dataset of reindeer calving dates recorded since 1970 in a semidomesticated reindeer population in Finnish Lapland and proxies of plant phenology; we tested the main hypothesis that the time lag between calving date and the plant phenology in autumn when females store nutrient reserves to finance reproduction would lead to consequences on reproductive success, as the time lag with spring conditions would. As predicted, the reproductive success of females of the Kutuharju reindeer population was affected by both the onset of spring green-up and vegetative senescence in autumn as calves were born heavier and with a higher first-summer survival when the onset of the vegetation growth was earlier and the end of the thermal growing season the previous year was earlier as well. Our results demonstrated that longer plant growing seasons might be detrimental to reindeer's reproductive success if a later end is accompanied by a reduced abundance of mushrooms.

Early-life conditions determine the between-individual heterogeneity in plasticity of calving date in reindeer.

Phenotypic plasticity has become a key concept to enhance our ability to understand the adaptive potential of species to track the pace of climate change by allowing a relatively rapid adjustment of life-history traits. Recently, population-level trends of an earlier timing of reproduction to climate change have been highlighted in many taxa, but only few studies have explicitly taken into consideration between-individual heterogeneity in phenotypic plasticity. Using a long-term data of a semi-domesticated reindeer (Rangifer tarandus) population, we demonstrated that females differed greatly in their mean calving date but only slightly in the magnitude of their plastic response to the amount of precipitation in April. We also showed that despite the absence of a population trend, females individually responded to the amount of precipitation in April by delaying their calving dates. Females' calving date under average climatic conditions was best predicted by their birthdate, by their physical condition in March-April-May before their first calving season and by their first calving date. The degree of their phenotypic plasticity was not dependent on any of the females' attributes early in life tested in this study. However, females who delayed their calving dates in response to a higher amount of precipitation in April slightly produced less calves over their reproductive life. These findings confirmed that early-life conditions of female reindeer can shape their phenotypic value during reproductive life, supporting the importance of maternal effects in shaping individuals' lifetime reproductive success. Whether females differed in the magnitude of their plastic response to climatic changes has received contrasted responses for various ungulate species. This calls for more research to enhance our understanding of the underlying mechanisms, leading to the complexity of plastic responses among populations to cope with current climate change.

Activity budget and spatial distribution of Bennett's wallabies (Macropus rufogriseus) in open versus closed exhibit designs.

Although many studies investigating the impacts of zoo exhibit designs on captive animals exist, none have been performed on how they influence the behavior and welfare of captive Bennett's wallabies (Macropus rufogriseus). Here, we assess the impact of exhibit design on the activity budget and spatial distribution of Bennett's wallabies. We compared animal behavior in two open exhibits (i.e. physical interaction between animals and visitors permitted) to two closed exhibits (i.e. physical interaction between animals and visitors prohibited). Behavioral data were collected using focal sampling, and spatial distribution was recorded on exhibit maps at regular time intervals. We found a significant increase in feeding and interactive behaviors in closed exhibits in comparison to open exhibits. However, other behaviors such as resting, locomotion, and vigilance did not vary with design. Functional use of space was similar between both designs; however, the effect of habituation may be relevant to consider in future studies. Although some support for visitor effects were present, our study provided no evidence for strong impacts of exhibit design on Bennett's wallaby welfare. Our study emphasizes the need for additional research into the impacts of how zoo environments affect Bennett's wallaby behavior and welfare.

Winter and spring climatic conditions influence timing and synchrony of calving in reindeer.

In a context of climate change, a mismatch has been shown to occur between some species' reproductive phenology and their environment. So far, few studies have either documented temporal trends in calving phenology or assessed which climatic variables influence the calving phenology in ungulate species, yet the phenology of ungulates' births affects offspring survival and population's recruitment rate. Using a long-term dataset (45 years) of birth dates of a semi-domesticated reindeer population in Kaamanen, North Finland, we show that calving season has advanced by ~ 7 days between 1970 and 2016. Advanced birth dates were associated with lower precipitation and a reduced snow cover in April and warmer temperatures in April-May. Improved females' physical condition in late gestation due to warmer temperatures in April-May and reduced snow conditions in April probably accounted for such advance in calving date. On the other hand, a lengthening of the calving season was reported following a warmer temperature in January, a higher number of days when mean temperature exceeds 0°C in October-November and a decreasing snow cover from October to November. By affecting the inter-individual heterogeneity in the plastic response of females' calving date to better climatic conditions in fall and winter, climatic variability contributed to weaken the calving synchrony in this herd. Whether variability in climatic conditions form environmental cues for the adaptation of calving phenology by females to climate change is however uncertain, but it is likely. As such this study enhances our understanding on how reproductive phenology of ungulate species would be affected by climate change.

Diagnosis, Prevalence, Awareness, Treatment, Prevention, and Control of Hypertension in Cameroon: Protocol for a Systematic Review and Meta-Analysis of Clinic-Based and Community-Based Studies.

BACKGROUND: Hypertension holds a unique place in population health and health care because it is the leading cause of cardiovascular disease and the most common noncommunicable condition seen in primary care worldwide. Without effective prevention and control, raised blood pressure significantly increases the risk of stroke, myocardial infarction, chronic kidney disease, heart failure, dementia, renal failure, and blindness. There is an urgent need for stakeholders-including individuals and families-across the health system, researchers, and decision makers to work collaboratively for improving prevention, screening and detection, diagnosis and evaluation, awareness, treatment and medication adherence, management, and control for people with or at high risk for hypertension. Meeting this need will help reduce the burden of hypertension-related disease, prevent complications, and reduce the need for hospitalization, costly interventions, and premature deaths. OBJECTIVE: This review aims to synthesize evidence on the epidemiological landscape and control of hypertension in Cameroon, and to identify elements that could potentially inform interventions to combat hypertension in this setting and elsewhere in sub-Saharan Africa. METHODS: The full search process will involve several steps, including selecting relevant databases, keywords, and Medical Subject Headings (MeSH); searching for relevant studies from the selected databases; searching OpenGrey and the Grey Literature Report for gray literature; hand searching in Google Scholar; and soliciting missed publications (if any) from relevant authors. We will select qualitative, quantitative, or mixed-methods studies with data on the epidemiology and control of hypertension in Cameroon. We will include published literature in French or English from electronic databases up to December 31, 2016, and involving adults aged 18 years or older. Both facility and population-based studies on hypertension will be included. Two reviewers of the team will independently search, screen, extract data, and assess the quality of selected studies using suitable tools. Selected studies will be analyzed by narrative synthesis, meta-analysis, or both, depending on the nature of the data retrieved in line with the review objectives. RESULTS: This review is part of an ongoing research program on disease prevention and control in the context of the dual burden of communicable and noncommunicable diseases in Africa. The first results are expected in 2017. CONCLUSIONS: This review will provide a comprehensive assessment of the burden of hypertension and control measures that have been designed and implemented in Cameroon. Findings will form the knowledge base relevant to stakeholders across the health system and researchers who are involved in hypertension prevention and control in the community and clinic settings in Cameroon, as a yardstick for similar African countries. TRIAL REGISTRATION: PROSPERO registration number: CRD42017054950; http://www.crd.york.ac.uk/PROSPERO/ display_record.asp?ID=CRD42017054950 (Archived by WebCite at http://www.webcitation.org/6qYSjt9Jc).

Temporal variation in the operational sex ratio and male mating behaviours in reindeer (Rangifer tarandus).

In polygynous species, sexual selection is mostly driven by male ability to monopolize access to females in oestrous. In ungulates, the operational sex ratio (OSR), i.e. the proportion of males to individuals ready to mate, varies throughout the peak rut, resulting from the temporal variation in the number of females in oestrous. But the way males adjust their mating tactics to maximise their access to females in oestrous (i.e. as OSR varies) is yet to be investigated. Using 15 years of behavioural observations in reindeer (Rangifer tarandus), we compared the relative importance of time within the rutting season (days to the peak-rut) and the OSR to explain the variation in the propensity (i.e. the frequency after controlling for the potential number of encounters) of young and adult dominant males to engage in four mating tactics: herding females, chasing other males, investigating female reproductive status, and courting females. Male-male agonistic behaviour was the most frequent mating behaviour, followed by herding. As predicted, dominant male mating tactics changed over the rutting season: first herding, then chasing other males, and finally investigating and courting females. In contrast to our prediction, we did not find support for the OSR theory. We noted some discrepancies in how young and adult dominant males adjusted their tactics during the mating season, adults being more efficient in timing and in performing their behaviour to maximise access to females in oestrous. The reported sequence of mating tactics may be more efficient than a static mating tactic to monopolize females in oestrous, regardless of the population composition.

Changes in the dominance hierarchy of captive female Japanese macaques as a consequence of merging two previously established groups.

Dominance hierarchies play an important role in reducing competition and aggression in social animals. In zoos, changes in group composition are often required due to management protocols, but these changes may have long lasting effects on dominance hierarchies, and, consequently, the wellbeing of the animals. We studied the changes in the female dominance hierarchy that occurred both during and after the formation of a group of 10 adult Japanese macaques at the Zoo de Granby by combining members from two previously established groups. There was no significant correlation between individual ranks in the old groups (groups A and B) and their ranks in the new group (group AB), indicating a significant change in the hierarchy. Alliances between kin appeared to be important in determining rank; when the sister of the dominant female was removed from group AB, the hierarchy changed significantly a second time. The average standardized rank of individuals added later in the formation process of group AB was not different from those added earlier. Ranks in the group AB did correlate with age of individual at the beginning of the field season, but not at the end, after the shift in hierarchy occurred. Zoo management must be aware of the consequences small changes in a social group can have when removing and transferring individuals in both primates and in other social species. Zoo Biol. 35:505-512, 2016. © 2016 Wiley Periodicals, Inc.

Foraging competition in larger groups overrides harassment avoidance benefits in female reindeer (Rangifer tarandus).

Male harassment toward females during the breeding season may have a negative effect on their reproductive success by disturbing their foraging activity, thereby inducing somatic costs. Accordingly, it is predicted that females will choose mates based on their ability to provide protection or will aggregate into large groups to dilute per capita harassment level. Conversely, increasing group size may also lead to a decrease in foraging activity by increasing foraging competition, but this effect has rarely been considered in mating tactic studies. This study examined the importance of two non-exclusive hypotheses in explaining the variations of the female activity budget during the breeding season: the male harassment hypothesis, and the female foraging competition hypothesis. We used focal observations of female activity from known mating groups collected during the breeding season from a long-term (15 years) study on reindeer Rangifer tarandus. We found that females were more disturbed (i.e., spent less time feeding) in the presence of young dominant males, and marginally disturbed in the presence of satellite males, which supports the male harassment hypothesis. We also found that female disturbance level increased with group size, being independent of the adult sex ratio. Consequently, these results rejected the dilution effect, but strongly supported the foraging competition hypothesis. This study therefore highlights a potential conflict in female behaviour. Indeed, any gains from harassment protection were negated by an increase of 6-7 females, since adult males lead larger groups than young males.

Allosuckling in reindeer (Rangifer tarandus): milk-theft, mismothering or kin selection?

Allosuckling, the suckling of offspring from females other than their own mother, has been reported in a number of mammalian species, including reindeer. The causes and function of this behaviour are still being investigated. We monitored 25 doe-calf pairs of semi-domestic reindeer Rangifer tarandus over 10 weeks to test three allosuckling/allonursing hypotheses: (1) milk theft, calves opportunistically allosuckle; (2) mismothering, misdirected maternal care; and (3) kin-selected allonursing. A calf soliciting an allosuckling bout was categorized as non-filial (NF), and a calf soliciting a suckling bout from its mother was categorized as filial (F). We recorded 9757 solicitations, of which 5176 were successful F bouts and 1389 were successful NF bouts. The rejection rates were greater for NF than F calves. The proportions of antiparallel positions adopted were greater for F than NF calves. The odds of an allobout were lower for calves arriving 1st, 3rd, 4th, 5th or 6th than for those arriving 2nd, but the odds did not vary with position adopted and relatedness. Our results provided support to the milk-theft hypothesis, whereas limited support for the mismothering hypothesis was found. Our results did not support the hypothesized kin selection function of allosuckling in reindeer. This article is part of a Special Issue entitled: insert SI title.

Highly competitive reindeer males control female behavior during the rut.

During the rut, female ungulates move among harems or territories, either to sample mates or to avoid harassment. Females may be herded by a male, may stay with a preferred male, or aggregate near a dominant male to avoid harassment from other males. In fission-fusion group dynamics, female movement is best described by the group's fission probability, instead of inter-harem movement. In this study, we tested whether male herding ability, female mate choice or harassment avoidance influence fission probability. We recorded group dynamics in a herd of reindeer Rangifer tarandus equipped with GPS collars with activity sensors. We found no evidence that the harassment level in the group affected fission probability, or that females sought high rank (i.e. highly competitive and hence successful) males. However, the behavior of high ranked males decreased fission probability. Male herding activity was synchronous with the decrease of fission probability observed during the rut. We concluded that male herding behavior stabilized groups, thereby increasing average group size and consequently the opportunity for sexual selection.

Age-specific changes in different components of reproductive output in female reindeer: terminal allocation or senescence?

Two different processes can lead to a change in individual reproductive output with age in long-lived iteroparous vertebrates. The senescence hypothesis predicts a decline of performance in old age, whereas the terminal allocation hypothesis predicts an increase. Using long-term (>30 years) individually based data of female reindeer, we first assessed age-specific variation in body mass and different components of reproductive output. Then we investigated the contribution of senescence and terminal allocation (the increase in components of reproductive output) processes for shaping observed patterns. We found that female reindeer body mass increased up to about 11.5 years of age, and decreased afterwards, supporting the senescence hypothesis. Calf birth mass, both in absolute terms or for a given female mass, first increased and then declined with female age, also supporting the senescence hypothesis. The female mass gain (June-September) decreased with increasing age, and female change in mass between 2 consecutive years decreased with female age, all patterns again supporting the senescence hypothesis. However, the autumn calf mass did not change with age. Calf body mass in autumn tended to be positively related to female mass gain, supporting a quality effect. Raising a calf had a marked negative effect on female mass gain, indicating energetic reproductive costs of raising a calf. Calf body mass in autumn did not influence yearly female mass change. Overall, our results provided consistent evidence for general effects of senescence on most components of reproductive output and highlighted that both individual heterogeneity and reproductive costs shape female reindeer reproductive tactics.

Feedback effects of chronic browsing on life-history traits of a large herbivore.

1. Increasing ungulate populations are affecting vegetation negatively in many areas, but few studies have assessed the long-term effects of overbrowsing on individual life-history traits of ungulates. 2. Using an insular population of white-tailed deer (Odocoileus virginianus Zimmermann; Anticosti, Québec, Canada) introduced in 1896, and whose density has remained high since the first evidence of severe browsing in the 1930s, we investigated potential feedbacks of long-term and heavy browsing on deer life-history traits. 3. We assessed whether chronic browsing contributed to a decline of the quality of deer diet in early autumn during the last 25 years, and evaluated the impacts of reduced diet quality on deer body condition and reproduction. 4. Rumen nitrogen content declined 22% between two time periods, 1977-79 and 2002-04, indicating a reduction in diet quality. 5. After accounting for the effects of year within the time period, age and date of harvest in autumn, peak body mass of both sexes declined between the two time periods. At the end of November, males were on average 12% heavier and adult does 6% heavier in 1977-79 than in 2002-04. Hind foot length did not vary between time periods. 6. The probability of conception increased 15% between the two time periods, but litter size at ovulation declined 7%, resulting in a similar total number of ovulations in 2002-04 and in 1977-79. 7. Our results suggest that following a decline in diet quality, white-tailed deer females modified their life-history strategies to maintain reproduction at the expense of growth. 8. Deer appear to tolerate drastic reductions in diet quality by modifying their life history traits, such as body mass and reproduction, before a reduction in density is observed. Such modifications may contribute to maintain high population density of large herbivores following population irruption.

Heterogeneity in individual quality overrides costs of reproduction in female reindeer.

Reproductive allocation at one age is predicted to reduce the probability of surviving to the next year or to lead to a decrease in future reproduction. This prediction assumes that reproduction involves fitness costs. However, few empirical studies have assessed whether such costs may vary with the age at primiparity or might be overridden by heterogeneities in individual quality. We used data from 35 years' monitoring of individually marked semi-domestic reindeer females to investigate fitness costs of reproduction. Using multi-state statistical models, we compared age-specific survival and reproduction among four reproductive states (never reproduced, experienced non-breeders, reproduced but did not wean offspring, and reproduced and weaned offspring) and among contrasted age at primiparity. We assessed whether reproductive costs occurred, resulting in a trade-off between current reproduction and future reproduction or survival, and whether early maturation was costly or rather reflected differences in individual quality of survival and reproduction capabilities. We did not find any evidence for fitness costs of reproduction in female reindeer. We found no cost of gestation and lactation in terms of future reproduction and survival. Conversely, successful breeders had higher survival and subsequent reproductive success than experienced non-breeders and unsuccessful breeders, independently of the age at primiparity. Moreover, it was beneficial to mature earlier, especially for females that successfully weaned their first offspring. Successful females at early primiparity remained successful throughout their life, clearly supporting the existence of marked among-female differences in quality. The weaning success peaked for multiparous females and was lower for first-time breeders, indicating a positive effect of experience on reproductive performance. Our findings emphasize an overwhelming importance of individual quality and experience to account for observed variation in survival and reproductive patterns of female reindeer that override trade-offs between current reproduction and future performance, at least in the absence of harsh winters.

No evidence of inbreeding avoidance in a polygynous ungulate: the reindeer (Rangifer tarandus).

In polygynous species, mate choice is an integrated part of sexual selection. However, whether mate choice occurs independently of the genetic relatedness among mating pairs has received little attention, although inbreeding may have fitness consequences. We studied whether genetic relatedness influenced females' choice of partner in a highly polygynous ungulate--the reindeer (Rangifer tarandus)--in an experimental herd during two consecutive rutting seasons; the herd consisting of 75 females in 1999 and 74 females in 2000 was exposed to three 4.5-year-old adults and three 1.5-year-old young males, respectively. The females' distribution during peak rut was not influenced by their genetic relatedness with the dominant males of the mating groups. Further, genetic relatedness did not influence the actual choice of mating partner. We conclude that inbreeding avoidance through mating group choice as well as choice of mating partner, two interconnected processes of female mate choice operating at two different scales in space and time, in such a highly female-biased reindeer populations with low level of inbreeding may not occur.

Good reindeer mothers live longer and become better in raising offspring.

Longevity is the main factor influencing individual fitness of long-lived, iteroparous species. Theories of life history evolution suggest this is because increased longevity allows individuals to (i) have more breeding attempts (time component), (ii) accumulate experience so as to become better able to rear offspring (experience component) or (iii) because individuals reaching old age have above-average quality (quality component). We assess empirically the relative influences of time, experience and quality on the relationship between longevity and individual fitness among female reindeer. Fitness increased with longevity due to all three processes. All females increased in success with age up to their penultimate year of life (experience component), the success of the terminal-breeding occasion was strongly dependent on longevity. Long-lived females had more successful breeding attempts during their life (time component), and had higher reproductive success at all ages, especially during the last year of life (individual quality component) than short-lived females. Our study reveals a more complex relationship between longevity and fitness in large mammals than the simple increase of the number of reproductive attempts when living longer.

Adaptive adjustment of offspring sex ratio and maternal reproductive effort in an iteroparous mammal.

Large mammals in seasonal environments have a pattern of high-reproductive synchrony in spring, but how the timing of reproduction affects resource allocation decisions at different stages of the reproductive cycle remains largely unexplored. By manipulating the timing of conception in reindeer (Rangifer tarandus), we tested how the timing of conception affected sex ratio, gestation length and weight development of mother and offspring. Females that conceived at their first ovulation within the rut had a 60.5% probability of producing a male; in contrast, females that conceived a cycle later had a 31.3% probability of producing a male. Late conceiving females had gestation times that were 10 days shorter and the calves were 0.6 kg (9.2%) lighter at birth and 7.4 kg (14.7%) lighter in autumn. Over the year, female weight changes was similar between the groups suggesting reindeer follow a bet-hedging strategy; reducing the quality of this year's offspring to ensure their own future reproduction and survival. Harvesting is often selective leading to skewed sex ratios and age structure, which may influence the timing of reproduction due to females hesitation to mate with young males. Whenever this hesitation is strong enough to increase the frequency of recycling, harvesting is likely to have profound life history consequences.