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Martin, Alexandre M.; Festa-Bianchet, Marco; Coltman, David W.; Pelletier, Fanie 2016-06-02 Sexual selection has a critical role in evolution, and it is fundamental to identify what ecological factors drive its variation. Disentangling the ecological correlates of sexual selection over the long term, however, is challenging and has rarely been done in nature. We sought to assess how demographic changes influenced the intensity, direction and form of sexual selection and whether selective pressures varied with age. We tested whether breeder sex ratio, number of competitors and age structure influenced selection differentials on horn length of wild bighorn rams (Ovis canadensis) of different age classes on Ram Mountain, Alberta. We used 21 years of data including a detailed pedigree, demographic parameters and repeated morphological measurements. Sexual selection on horn length of males of all ages was directional and positive. Selection intensity increased with the number of competitors, reflecting male–male encounter rate during the rut, but was independent of breeder sex ratio or age structure. This result can also be linked to changes in population size because the number of competitors was highly correlated to total number of sheep. This demographic effect likely arises from age-dependent mating tactics. Males aged 2–4 years are weakly competitive and experienced stronger sexual selection as they accounted for a greater proportion of all males. Selection experienced by mature males appeared independent of demography. Our study provides a rare description of the demographic determinants of sexual selection in nature.
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Hamel, Sandra; Gaillard, Jean-Michel; Douhard, Mathieu; Festa-Bianchet, Marco; Pelletier, Fanie; Yoccoz, Nigel G. 2017-09-21 Heterogeneity among individuals influences the life-history trajectories we observe at the population level because viability selection, selective immigration and emigration processes, and ontogeny change the proportion of individuals with specific trait values with increasing age. Here, we review the two main approaches that have been proposed to account for these processes in life-history trajectories, contrasting how they quantify ontogeny and selection, and proposing ways to overcome some of their limitations. Nearly all existing approaches to model individual heterogeneity assume either a single normal distribution or a priori known groups of individuals. Ontogenetic processes, however, can vary across individuals through variation in life-history tactics. We show the usefulness of describing ontogenetic processes by modelling trajectories with a mixture model that focuses on heterogeneity in life-history tactics. Additionally, most methods examine individual heterogeneity in a single trait, ignoring potential correlations among multiple traits caused by latent common sources of individual heterogeneity. We illustrate the value of using a joint modelling approach to assess the presence of a shared latent correlation and its influence on life-history trajectories. We contrast the strengths and limitations of different methods for different research questions, and we exemplify the differences among methods using empirical data from long-term studies of ungulates.
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Poisson, Yoanna; Festa-Bianchet, Marco; Pelletier, Fanie 2019-12-12 <ol> <li style="text-align:justify;margin-bottom:11px;">Sustainable exploitation must minimize its impact on the ecology and evolution of exploited wildlife. Intense phenotype-based selective harvests can induce evolutionary change. Refuges could mitigate those evolutionary effects if individuals not subject to selective hunting in harvest refuges migrated and reproduced in hunted areas. The role of harvest refuges on phenotypic rescue of trophy-hunted species, however, has rarely been tested.</li> <li style="text-align:justify;margin-bottom:11px;">We investigated spatial and temporal variation in the effect of refuges on horn size and age at harvest in bighorn sheep (<i>Ovis canadensis</i>). We analyzed data on 5 826 males harvested over 39 years in Alberta, Canada.</li> <li style="text-align:justify;margin-bottom:11px;">Horn length, a trait targeted by hunters, and age at harvest increased with the amount of protected areas 5-40 km around each kill. Horn base circumference, however, was independent of proximity to refuges.</li> <li style="text-align:justify;margin-bottom:11px;">The number of males harvested increased during the last 10 days of the hunting season in late October, corresponding with the timing of bighorn male breeding migrations. Males shot during those 10 days were on average 17% closer to a refuge than males shot earlier in the season. Apparently, some large males exit refuges late in the hunting season, are shot, and cannot contribute to rescue. Uncertainty remains about the proportion of males exiting refuges after the hunting season and how many survive to reproduce.</li> <li style="text-align:justify;margin-bottom:11px;"> <i>Synthesis and applications</i>. Harvest refuges are unlikely to rescue hunted populations of bighorn sheep in Alberta, because some males exiting refuges are at risk of harvest before they mate. For phenotypic rescue to be effective, unselected males must reproduce before they are shot. Closing the hunting season 10 days earlier would increase survival of unselected rams exiting refuges.</li> </ol>
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Martin, Alexandre M.; Festa-Bianchet, Marco; Coltman, David W.; Pelletier, Fanie 2014-11-25 The reliability and consistency of the many measures proposed to quantify sexual selection have been questioned for decades. Realized selection on quantitative characters measured by the selection differential i was approximated by metrics based on variance in breeding success, using either the opportunity for sexual selection Is or indices of inequality. There is no consensus about which metric best approximates realized selection on sexual characters. Recently, the opportunity for selection on character mean OSM was proposed to quantify the maximum potential selection on characters. Using 21 years of data on bighorn sheep (Ovis canadensis), we investigated the correlations between seven indices of inequality, Is, OSM and i on horn length of males. Bighorn sheep are ideal for this comparison because they are highly polygynous, sexually dimorphic, ram horn length is under strong sexual selection, and we have detailed knowledge of individual breeding success. Different metrics provided conflicting information, potentially leading to spurious conclusions about selection patterns. Iδ, an index of breeding inequality, and to a lesser extent Is, showed the highest correlation with i on horn length, suggesting that these indices document breeding inequality in a selection context. OSM on horn length was strongly correlated with i, Is, and indices of inequality. By integrating information on both realized sexual selection and breeding inequality, OSM appeared to be the best proxy of sexual selection and may be best suited to explore its ecological bases.
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Ritchot, Yanny; Pelletier, Fanie; Festa-Bianchet, Marco; Coltman, David 2022-03-26 <p style="margin-bottom:11px;">In long-lived polygynous species, male reproductive success is often monopolized by a few mature dominant individuals. Young males are generally too small to be dominant and may employ alternative tactics, however, little is known about the determinants of reproductive success for young males. Understanding the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long-term trade-offs among life-history traits. Selective pressures driven by fluctuating environmental conditions may depend on age-class. We evaluated the determinants of reproduction in male bighorn sheep (<em>Ovis canadensis</em>) aged 2-4 years using 30 years of individual-level data. These young males cannot defend estrous ewes and use alternative mating tactics. We also investigated how the age of first detected reproduction was correlated to lifetime reproductive success and longevity. We found that reproductive success of males aged three years was positively correlated to body mass, to the proportion of males aged 2-4 years in the competitor pool and to the number of females available per adult male. These results suggest that reproductive success depends on both competitive ability and population age-sex structure. None of these variables, however, had significant effects on the reproductive success of males aged 2 or 4 years. Known reproduction before the age of five increased lifetime reproductive success but decreased longevity, suggesting a long-term survival cost of early reproduction. Our analyses reveal that both individual-level phenotypic and population-level demographic variables influence reproductive success by young males and provide a rare assessment of fitness trade-offs in wild polygynous males.</p>
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Poirier, Marc-Antoine; Coltman, David W.; Pelletier, Fanie; Jorgenson, Jon; Festa-Bianchet, Marco 2018-08-31 Isolation of small populations is expected to reduce fitness through inbreeding and loss of genetic variation, impeding population growth and compromising population persistence. Species with long generation time are the least likely to be rescued by evolution alone. Management interventions that maintain or restore genetic variation to assure population viability are consequently of significant importance. We investigated, over 27 years, the genetic and demographic consequences of a demographic bottleneck followed by artificial supplementation in an isolated population of bighorn sheep (Ovis canadensis). Based on a long-term pedigree and individual monitoring, we documented the genetic decline, restoration and rescue of the population. Microsatellite analyses revealed that the demographic bottleneck reduced expected heterozygosity and allelic diversity by 6.2 and 11.3%, respectively, over two generations. Following supplementation, first-generation admixed lambs were 6.4% heavier at weaning and had 28.3% higher survival to 1 year compared to lambs of endemic ancestry. Expected heterozygosity and allelic diversity increased by 4.6 and 14.3% after two generations through new alleles contributed by translocated individuals. We found no evidence for outbreeding depression and did not see immediate evidence of swamping of local genes. Rapid intervention following the demographic bottleneck allowed the genetic restoration and rescue of this bighorn sheep population, likely preventing further losses at both the genetic and demographic levels. Our results provide further empirical evidence that translocation can be used to reduce inbreeding depression in nature and has the potential to mitigate the effect of human-driven environmental changes on wild population.
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Gélin, Uriel; Wilson, Michelle E.; Coulson, Graeme; Festa-Bianchet, Marco 2014-07-04 1. When resources are scarce, female mammals should face a trade-off between lactation and other life-history traits such as growth, survival and subsequent reproduction. Kangaroos are ideal to test predictions about reproductive costs because they may simultaneously lactate and carry a young, and have indeterminate growth and a long breeding season. 2. An earlier study in three of our five study populations prevented female eastern grey kangaroos (Macropus giganteus) from reproducing during one reproductive season by either inserting contraceptive implants or removing very small pouch young. We explored how individual and environmental variables affect the costs of reproduction over time, combining this experimental reduction of reproductive effort with multi-year monitoring of 270 marked females. Experimental manipulation should control for individual heterogeneity, revealing the costs of reproduction and their likely sources. We also examined the fitness consequences of reproductive effort and offspring sex among unmanipulated individuals to test if sex allocation strategies affected trade-offs. 3. Costs of reproduction included longer inter-birth intervals and lower probability of producing a young that survived to 7 months in the subsequent reproductive event. Weaning success, however, did not differ significantly between manipulated and control females. By reducing reproductive effort, manipulation appeared to increase individual condition and subsequent reproductive success. 4. Effects of offspring sex upon subsequent reproductive success varied according to year and study population. Mothers of sons were generally more likely to have a young that survived to 7 months, compared to mothers of daughters. 5. The fitness costs of reproduction arise from constraints in both acquisition and allocation of resources. To meet these costs, females delay subsequent parturition and may manipulate offspring sex. Reproductive tactics thus vary according to the amount of resource available to each individual, promoting a wide range in reproductive performance within and among individuals and populations.
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Renaud, Limoilou-Amelie; Festa-Bianchet, Marco; Pelletier, Fanie 2021-10-21 <p style="text-align:start;text-indent:0px;"><span><span style="font-style:normal;"><span><span style="font-weight:normal;"><span style="letter-spacing:normal;"><span><span><span style="white-space:normal;"><span><span><span>In species with long gestation, females commit to reproduction several months before parturition. If cues driving conception date are uncoupled from spring conditions, parturition could be mistimed. Mismatch may increase with global change if the rate of temporal changes in autumn cues differs from the rate of change in spring conditions. Using 17 years of data on climate and vegetation phenology, we show that autumn temperature and precipitation, but not vegetation phenology, explain parturition date in bighorn sheep. Although autumn cues drive the timing of conception, they do not predict conditions at parturition in spring. We calculated the mismatch between individual parturition date and spring green-up, assessed whether mismatch increased over time and investigated the consequences of mismatch on lamb neonatal survival, weaning mass and overwinter survival. Mismatch fluctuated over time but showed no temporal trend. Temporal changes in green-up date did not lead to major fitness consequence of mismatch. Detailed data on individually marked animals revealed no effect of mismatch on neonatal or overwinter survival, but lamb weaning mass was negatively affected by mismatch. Capital breeders might be less sensitive to mismatch than income breeders because they are less dependent on daily food acquisition. Herbivores in seasonal environments may access sufficient forage to sustain lactation before or after the spring ‘peak’ green-up, and partly mitigate the consequences of a mismatch. Thus, the effect of phenological mismatch on fitness may be affected by species life-history, highlighting the complexity in quantifying trophic mismatches in the context of climate change. </span></span></span></span></span></span></span></span></span></span></span></p>
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Douhard, Mathieu; Festa-Bianchet, Marco; Coltman, Dave W.; Pelletier, Fanie; Coltman, David W. 2016-01-13 Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex-ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex-biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.
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Douhard, Mathieu; Pigeon, Gabriel; Festa-Bianchet, Marco; Coltmann, Dave W.; Guillemette, Simon; Pelletier, Fanie; Coltman, David W. 2016-11-30 The development of male secondary sexual characters such as antlers or horns has substantial biological and socio-economic importance because in many species these traits affect male fitness positively through sexual selection and negatively through trophy hunting. Both environmental conditions and selective hunting can affect horn growth but their relative importance remains unexplored. We first examined how a large-scale climate index, the Pacific Decadal Oscillation (PDO), local weather and population density influenced both absolute and relative annual horn growth from birth to 3 years of male bighorn sheep (Ovis canadensis) over 42 years. We then examined the relative influence of environmental conditions and evolution mainly driven by trophy hunting on male horn length at 3 years of age. Horn growth was positively influenced by low population density and warm spring temperature, suggesting that ongoing climate change should lead to larger horns. Seasonal values of PDO were highly correlated. Horn growth increased with PDO in spring or summer at low density, but was weak at high density regardless of PDO. The interaction between population density and PDO in spring or summer accounted for a similar proportion of the observed annual variation in horn growth (32% or 37%) as did the additive effects of spring temperature and density (34%). When environmental conditions deteriorated, males allocated relatively more resources to summer mass gain than to horn growth, suggesting a conservative strategy favoring maintenance of condition over allocation to secondary sexual characters. Population density explained 27% of the variation in horn length, while evolutionary effects explained 9% of the variance. Thus, our study underlines the importance of both evolution and phenotypic plasticity on the development of a secondary sexual trait.
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Miller, Joshua M.; Malenfant, René M.; David, Patrice; Davis, Corey S.; Poissant, Jocelyn; Hogg, John T.; Festa-Bianchet, Marco; Coltman, David W. 2013-09-17 Heterozygosity–fitness correlations (HFCs) are often used to link individual genetic variation to differences in fitness. However, most studies examining HFCs find weak or no correlations. Here, we derive broad theoretical predictions about how many loci are needed to adequately measure genomic heterozygosity assuming different levels of identity disequilibrium (ID), a proxy for inbreeding. We then evaluate the expected ability to detect HFCs using an empirical data set of 200 microsatellites and 412 single nucleotide polymorphisms (SNPs) genotyped in two populations of bighorn sheep (Ovis canadensis), with different demographic histories. In both populations, heterozygosity was significantly correlated across marker types, although the strength of the correlation was weaker in a native population compared with one founded via translocation and later supplemented with additional individuals. Despite being bi-allelic, SNPs had similar correlations to genome-wide heterozygosity as microsatellites in both populations. For both marker types, this association became stronger and less variable as more markers were considered. Both populations had significant levels of ID; however, estimates were an order of magnitude lower in the native population. As with heterozygosity, SNPs performed similarly to microsatellites, and precision and accuracy of the estimates of ID increased as more loci were considered. Although dependent on the demographic history of the population considered, these results illustrate that genome-wide heterozygosity, and therefore HFCs, are best measured by a large number of markers, a feat now more realistically accomplished with SNPs than microsatellites.
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Pelletier, Fanie; Festa-Bianchet, Marco; Jorgenson, Jon T.; Feder, Chiarastella; Hubbs, Anne 2014-08-14 Human harvests can unwittingly drive evolution on morphology and life history, and these selective effects may be detrimental to the management of natural resources. Although theory suggests that harvest refuges, as sources of unselected animals, could buffer the effects of human exploitation on wild populations, few studies have assessed their efficiency. We analyzed records from >7000 trophy bighorn rams (Ovis canadensis) harvested in Alberta, Canada, between 1974 and 2011 to investigate if the movement of rams from refuges toward harvested areas reduced the effects of selective harvesting on horn size through phenotypic rescue. Rams taken near refuges had horns on average about 3% longer than rams shot far from refuges and were slightly older, suggesting migration from refuges into hunted areas. Rams from areas adjacent to and far from harvest refuges, however, showed similar declines in horn length and increases in age at harvest over time, indicating a decreasing rate of horn growth. Our study suggests that the influx of rams from refuges is not sufficient to mitigate the selective effects of sheep trophy harvest. Instead, we suggest that selective hunting of highly mobile animals may affect the genetic structure of populations that spend part of the year inside protected areas.
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Martin, Alexandre M.; Festa-Bianchet, Marco; Coltman, David W.; Pelletier, Fanie 2014-02-17 The evolution of conspicuous sexually selected traits, such as horns or antlers, has fascinated biologists for more than a century. Elaborate traits can only evolve if they substantially increase reproduction, because they probably incur survival costs to the bearer. Total selection on these traits, however, includes sexual selection on sires and viability selection on offspring and can be influenced by changes in each of these components. Non-random associations between paternal phenotype and offspring viability may thus affect total selection on sexually selected traits. Long-term data on wild bighorn sheep (Ovis canadensis) provide the first evidence in nature that association between paternal phenotype and lamb viability strengthens total selection on horn size of adult rams, a sexually selected trait. The association of paternal horn length and offspring viability was sexually antagonistic: long-horned males sired sons with high viability but daughters of low viability. These results shed new light on the evolutionary dynamics of an iconic sexually selected trait and have important implications for sustainable wildlife management.
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Montana, Luca; Rousseu, François; Garant, Dany; Festa-Bianchet, Marco 2020-07-28 In polygynous species, male reproductive success is predicted to be monopolized by a few dominant males. This prediction is often not supported, suggesting that ecological and alternative mating tactics influence siring success. The spatiotemporal distribution of individuals and the number of males competing for each receptive female are often overlooked, because they are difficult to monitor in wild animals. We examined how spatial overlap of female-male pairs, the time spent by a male on the breeding site, number of competitors and morphological traits influence siring probability in eastern grey kangaroos (<i>Macropus giganteus</i>). We compared home range overlap for 12,208 dam-male pairs and 295 known dam-sire pairs to define local competitive groups and to estimate every male’s opportunity to sire the young of each female. We compared models considering morphological traits relative to the entire population or to local competitive groups. Including local competition improved model performance because it estimated the intensity of competition and compared each male’s morphological traits to those of its competitive group. Regardless of size, males can increase their probability to sire a young by increasing their <i>mating opportunity</i> relative to the mother. We underline the importance of considering spatial structure to assess the intensity of competition in species where males cannot equally access all females in a population. The estimation of mating opportunity and intensity of local competition improves our understanding of how morphological traits affect siring success when each mating event involves a different set of competing males, a characteristic of most wild species.
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Douhard, Mathieu; Festa-Bianchet, Marco; Landes, Julie; Pelletier, Fanie 2019-03-07 1. Environmental conditions during early development, from conception to sexual maturity, can have lasting consequences on fitness components. Although adult lifespan often accounts for much of the variation in fitness in long-lived animals, we know little about how early environment affects adult lifespan in the wild, and even less about whether these effects differ between the sexes. 2. Using data collected over 45 years from wild bighorn sheep (Ovis canadensis), we investigated the effects of early environment on adult mortality in both sexes, distinguishing between natural and anthropogenic sources of mortality. 3. We used the average body mass of yearlings (at about 15 months of age) as a yearly index of environmental quality. We first examined sex differences in natural mortality responses to early environment by censoring harvested males in the year they were shot. We then investigated sex differences in the effects of early environment on overall mortality (natural and hunting mortality combined). Finally, we used path analysis to separate the direct influence of early environment from indirect influences, mediated by age at first reproduction, adult mass and horn length. 4. As early environmental conditions improved, natural adult mortality decreased in both sexes, although for males the effect was not statistically supported. Sex differences in the effects of early environment on adult mortality were detected only when natural and hunting mortality were pooled. Males that experienced favorable early environment had longer horns as adults and died earlier because of trophy hunting, which does not mimic natural mortality. Females that experienced favorable early environment started to reproduce earlier and early primiparity was associated with reduced mortality, suggesting a silver-spoon effect. 5. Our results show that early conditions affect males and females differently because of trophy hunting. These findings highlight the importance of considering natural and anthropogenic environmental factors across different life stages to understand sex differences in mortality.
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Toni, Pauline; Forsyth, David; Festa-Bianchet, Marco 2021-06-18 When the fitness costs and benefits of sons and daughters differ, offspring sex ratio manipulation could be an important reproductive tactic. We explored the effects of environment and maternal caring ability on offspring sex to test four adaptive sex ratio modification hypotheses: the Extrinsic Modification (EMH), Carrying Capacity (CCH), Trivers-Willard (TWH) and Cost-of-Reproduction (CRH) hypotheses. The EMH and CCH propose that environmental conditions shape offspring sex ratios, directly or in interaction with maternal condition. The TWH and CRH predict a positive relationship between maternal condition and production of the costlier sex. The TWH predicts that mothers with superior caring ability should produce more of the sex that can provide the greatest fitness returns from additional maternal allocation, and the CRH proposes that females with limited caring ability should reduce fitness costs by producing the cheaper sex. Repeated measures on 83 known-age eastern grey kangaroos, polygynous marsupials with strong sexual dimorphism, revealed that offspring sex ratio was independent of per capita forage, supporting neither the EMH nor CCH, but was dependent on maternal mass, consistent with the TWH and CCH. Our results, however, cannot clearly identify the ultimate cause of the relationship between maternal mass and greater production of sons. One of the three assumptions of the TWH could not be verified, and mothers of sons suffered only marginal additional fitness costs. Sex ratios in higher vertebrates are likely not solely explained by factors dependent on maternal control.
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Vuillaume, Barbara; Richard, Julien H.; hamel, sandra; Taillon, Joëlle; Festa-Bianchet, Marco; Côté, Steeve D. 2023 Analysis of the correlation between meteorological data from the North American Regional Reanalysis (NARR) and data available at Nunavik weather stations.Dataset used to produce the analysis for the electronic supplementary material 2, for the paper:Vuillaume, B., J. H. Richard, S. Hamel, J. Taillon, M. Festa-Bianchet and S. D. Côté. 2023. Birth date determines early calf survival in migratory caribou. https://creativecommons.org/licenses/by/4.0/legalcode
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Vuillaume, Barbara; Richard, Julien H.; hamel, sandra; Taillon, Joëlle; Festa-Bianchet, Marco; Côté, Steeve D. 2023 Dataset used to produce analyses of pre-weaning calf survival in migratory caribou, as part of the results presented in the core of the paper:Vuillaume, B., J. H. Richard, S. Hamel, J. Taillon, M. Festa-Bianchet and S. D. Côté. 2023. Birth date determines early calf survival in migratory caribou.2 datasets: including stillborns (..._withSB.txt) and excluding stillborns (..._noSB.txt)<br> https://creativecommons.org/licenses/by/4.0/legalcode
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Vuillaume, Barbara; Richard, Julien H.; hamel, sandra; Taillon, Joëlle; Festa-Bianchet, Marco; Côté, Steeve D. 2023 Dataset used for Fig. 2.A in the core of the paper :Vuillaume, B., J. H. Richard, S. Hamel, J. Taillon, M. Festa-Bianchet and S. D. Côté. 2023. Birth date determines early calf survival in migratory caribou.Also used in the supplementary material 3 of the article: Comparison of survival probabilities and environmental conditions encountered by female migratory caribou and their calves of the Rivière-aux-Feuilles herd (Nunavik, Canada) during the three years of the study (2016 – 2018). https://creativecommons.org/licenses/by/4.0/legalcode
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Vuillaume, Barbara; Richard, Julien H.; hamel, sandra; Taillon, Joëlle; Festa-Bianchet, Marco; Côté, Steeve D. 2023 Dataset used to select variables to test in CMR models as potential factors in pre-weaning calf survival in migratory caribou, in the paper:Vuillaume, B., J. H. Richard, S. Hamel, J. Taillon, M. Festa-Bianchet and S. D. Côté. 2023. Birth date determines early calf survival in migratory caribou. https://creativecommons.org/licenses/by/4.0/legalcode

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