Long‐lived and heavier females give birth earlier in roe deer

In seasonal environments, parturition of most vertebrates generally occurs within a short time‐window each year. This synchrony is generally interpreted as being adaptive, as early born young survive better over the critical season than late born young. Among large herbivores, the factors involved in driving among‐ and within‐individual variation in parturition date are poorly understood. We explored this question by analyzing the relative importance of attributes linked to female quality (longevity, median adult body mass and cohort), time‐dependent attributes linked to female condition (reproductive success the previous year, relative annual body mass and offspring cohort (year)), and age in shaping observed variation in parturition date of roe deer. A measure of quality combining the effects of female longevity and median adult body mass accounted for 11% of the observed among‐individual variation in parturition date. Females of 2 yr old give birth 5 d later than older females. Our study demonstrates that high quality (heavy and long‐lived) females give birth earlier than low quality females. Temporally variable attributes linked to female condition, such as reproductive success in the previous year and relative annual body mass, had no detectable influence on parturition date. We conclude that parturition date, a crucial determinant of reproductive success, is shaped by attributes linked to female quality rather than by time‐dependent attributes linked to female condition in income breeders (individuals that rely on current resource intake rather than on accumulated body reserves to offset the increased energy requirements due to reproduction) such as roe deer.     

Phenology and synchrony of roe deer breeding in northern Italy

The timing and synchrony of births of roe deerCapreolus capreolus Linnaeus, 1758 population were examined for 7 consecutive years in northern Italy. From 1997 to 2003, 136 fawns, 65 females and 71 males, from 117 litters were captured. The average date of fawning (30th of May) was constant among years and between sexes. The onset of parturition was 19th of May while the total length of the parturition season was 25 days (from 19th of May to 13th of June). Births were highly synchronous, with 80% of births occurring in 17 days. Synchrony of birth did not exhibit any significant interannual or intersexual variation. The stability of the birth pattern was confirmed by the parturition date of 13 radiocollared females who all gave birth within 4 days in consecutive fawning seasons. In agreement with previous findings, our data support the hypothesis that the roe deer is constrained at both ends of the fawning season by the short time between the fawning season and rut and by the need to match optimal climate conditions and forage quality to face late gestation and lactation. All these factors result in a relatively stable and synchronous birth season.     

Mismatch Between Birth Date and Vegetation Phenology Slows the Demography of Roe Deer

Marked impacts of climate change on biodiversity have frequently been demonstrated, including temperature-related shifts in phenology and life-history traits. One potential major impact of climate change is the modification of synchronization between the phenology of different trophic levels. High phenotypic plasticity in laying date has allowed many bird species to track the increasingly early springs resulting from recent environmental change, but although changes in the timing of reproduction have been well studied in birds, these questions have only recently been addressed in mammals. To track peak resource availability, large herbivores like roe deer, with a widespread distribution across Europe, should also modify their life-history schedule in response to changes in vegetation phenology over time. In this study, we analysed the influence of climate change on the timing of roe deer births and the consequences for population demography and individual fitness. Our study provides a rare quantification of the demographic costs associated with the failure of a species to modify its phenology in response to a changing world. Given these fitness costs, the lack of response of roe deer birth dates to match the increasingly earlier onset of spring is in stark contrast with the marked phenotypic responses to climate change reported in many other mammals. We suggest that the lack of phenotypic plasticity in birth timing in roe deer is linked to its inability to track environmental cues of variation in resource availability for the timing of parturition.     

Effects of cohort,sex, and birth date on body development of roe deer (Capreolus capreolus) fawns

We investigated the effects of the year of birth (cohort effect), sex and birth date on the variations observed in birth weight and postnatal growth rate in 209 newborn fawns marked during eight consecutive fawning seasons from a highly productive roe deer population under good nutritional conditions. Roe deer exhibited a fast body development with a birth weight of 1628 g and a postnatal growth rate of 139 g/day constant over the first 20 days of their life. As previously reported for dimorphic and polygynous ungulates, there was a marked cohort effect on the rate of body development. This variability was partly explained by climatic conditions during late gestation. When fewer than 5 days with rainfall over 5 mm occurred in April, fawns were heavier. High temperatures during April and during the winter could also be involved in fast body development of roe deer fawns. Sex of fawn did not affect roe deer growth pattern. This was expected on the basis of low sexual dimorphism in size and low polygyny level characteristic of roe deer. Lastly, the date of birth did not affect the body development of roe deer fawns. High constancy of mean birth dates and high synchrony of births observed in this population could account for this result.     

High red deer density depresses body mass of roe deer fawns

Many previous studies have pointed out that, when resources are limited, the potential for competition should be high among sympatric species that display overlaps in habitat and nutritional niches. However, reliable evidence of competition between red deer (Cervus elaphus) and roe deer (Capreolus capreolus) has not been yet reported for life history traits directly measuring performance such as body mass, reproduction, or survival. From long-term monitoring of deer populations in the reserve of La Petite Pierre (France), we measured the sex-specific responses of roe deer fawn body mass to changes in red deer density after accounting for possible confounding effects of date of shooting, climatic conditions, and roe deer density. As expected under the hypothesis of competition, red deer density in a given year had a marked negative influence on body mass of roe deer fawns born the same year and the following year. Fawn mass of roe deer males and females responded in similar ways to changes in red deer density. Our study provides the first evidence of a negative response of roe deer performance to high red deer density.     

Genetic variability and differentiation in Spanish roe deer (Capreolus capreolus): A phylogeographic reassessment within the European framework

Here we study 109 Iberian roe deer individuals corresponding to 9 Spanish populations. Individuals were sampled at locations that are expected to have acted as refugia for the species during the 20th century. Samples were analyzed for a 436 bp mtDNA fragment from the HVI region of mitochondrial DNA and 10 microsatellites. The 109 sequences gave 31 different haplotypes that enabled identification of a new haplogroup (mainly present in Northwestern Iberian populations and representing roughly a third of our samples) that is not present in other European roe deer populations. Using microsatellites, correspondence analysis and molecular coancestry information revealed high molecular differentiation among Northwestern and Central-Southern Spain roe deer populations. Both sequence and microsatellite analysis reveal that the Spanish roe deer populations are genetically heterogeneous and have high genetic structure clearly separating the Pyrenean-reintroduced populations and two main areas for the species in Spain (Northwestern and Central-Southern) coinciding with the two main areas acting as refugia for the majority of the mammal species during glaciations. The implications of the obtained information as regards the phylogeography of the species are discussed together with suggestions as to appropriate strategies for the conservation and management of populations.     

Advancing breeding phenology in response to environmental change in a wild red deer population

Most evidence for advances in phenology of in response to recent climate warming in wild vertebrate populations has come from long‐term studies of birds. Few studies have either documented phenological advances or tested their climatic causes and demographic consequences in wild mammal systems. Using a long‐term study of red deer on the Isle of Rum, Scotland, we present evidence of significant temporal trends in six phenological traits: oestrus date and parturition date in females, and antler cast date, antler clean date, rut start date and rut end date in males. These traits advanced by between 5 and 12 days across a 28‐year study period. Local climate measures associated with plant growth in spring and summer (growing degree days) increased significantly over time and explained a significant amount of variation in all six phenological traits, largely accounting for temporal advances observed in some of the traits. However, there was no evidence for temporal changes in key female reproductive performance traits (offspring birth weight and offspring survival) in this population, despite significant relationships between these traits and female phenology. In males, average antler weights increased over time presumably as a result of improved resource availability and physiological condition through spring and summer. There was no evidence for any temporal change in average male annual breeding success, as might be expected if the timing of male rutting behaviour was failing to track advances in the timing of oestrus in females. Our results provide rare evidence linking phenological advances to climate warming in a wild mammal and highlight the potential complexity of relationships between climate warming, phenology and demography in wild vertebrates.     

Costs of parturition and rearing in female sika deer (Cervus nippon)

The costs of parturition and lactation of female sika deer on Kinkazan Island (9.6 km(2) in size), northern Japan, which live at a high density (about 50 deer/km(2)), were evaluated by comparison of body weights of 481 females measured during a 15 year study (1993-2008). Weight data were chosen from only females that did not give birth in the preceding year. The mean body weight of females that did not give birth ("yelds") was significantly lower (P < 0.001) than that of females who gave birth ("milks"); yelds' body weight was 93.1% and 83.5% that of milks in the preceding and parturition years, respectively. The yelds increased in body weight by the following March by 8.2% (P < 0.001), whereas milks did not. Among the milks, those whose fawns survived until the following May ("rearing milks") lost body weight by 14.9% (P < 0.001). Milks who lost fawns within a week after birth ("early fawn-less milks") did not lose body weight (P = 0.583), while those whose fawns died after the first autumn but died before May ("late fawn-less milks") lost body weight by 19.9% (P < 0.001). These results indicate that sika deer females do not enter estrus unless they are heavy enough, and that both parturition and rearing are costly for sika deer mothers living in high-density conditions.     

Experimental evidence for density-dependence of home-range size in roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis> L.): a comparison of two long-term studies

The effect of experimental manipulation of population density on home-range size was investigated in two free-ranging roe deer (Capreolus capreolus) populations under contrasting environmental conditions. In these two long-term monitoring studies, one in Bogesund, Sweden (12 years) and one in Dourdan, France (10 years), deer density varied fourfold through varying culling pressure. Home-range data were collected by radio-tracking across the periods of contrasting density of the studies. We predicted that home-range size for females should vary in relation to the level of feeding competition, while for males, competition for mating opportunities should also influence range size, at least in summer when roe bucks are territorial. We found a highly consistent pattern over the two populations, with strong effects of deer density on home-range size, as well as significant differences between winter and summer ranges and between the sexes. Home ranges were consistently smaller at high density compared to low density. Males had larger ranges than females and this was particularly so during summer. Lastly, winter ranges were generally larger than summer ranges, particularly among females, although males at Dourdan had larger summer ranges compared to winter ranges. We suggest that the reduction of range size at high deer density during winter, as well as summer, is linked to the solitary behaviour and territorial social system of roe deer, with possible effects of dominance rank, even outside the mating season.     

Variation in cohort mandible size as an index of roe deer (Capredus capreolus) densities and population trends

This paper examines the effects of increasing population density on the skeletal development of roe deer as indexed by variation in the size of the mandible. We tested for density-dependence in total jaw length and minimum diastema height among cohorts in a single, intensively studied population in France. A strong relationship of average adult jaw length of both males and females with deer density in the year of birth of a given cohort was observed, and presumably reflects inadequate nutrition of juveniles at high population density. No such relationship was observed for the minimum diastema height which remained constant over the range of densities studied. This relationship was exploited to investigate population trends of roe deer in Britain. In nine populations, where equivalent data were available for between 16 and 28 separate cohorts, a consistent pattern of variation in cohort jaw size was observed, with a decrease in average total length, but not diastema height, towards the present. This trend is discussed in relation to changes in habitat structure in the coniferous forests of Britain. The utility of cohort jaw length as a management tool to index population trends of roe deer is highlighted.     

Age determination in roe deer — a new approach to tooth wear evaluated on known age in dividuals

A novel, simple, and objective method is presented for ageing roe deerCapreolus capreolus (Linnaeus, 1758) evaluated on 471 lower jaws from roe deer of known age (351 with permanent premolars). It is based on tooth eruption patterns and presence/absence of wear characters in jaws from roe deer integrated in a scoring system. Permanent cheek teeth emerge in May–July in the year af ter birth, which enables precise age determination of individuals with deciduous premolars. For individuals with permanent cheek teeth, the method provides the correct age for all individ uals younger than 13 months and > 80% of all individuals between 13 and 24 months old. For older in dividuals the accuracy decreases, but decent accuracy is achieved to the age of 48 months. Males have higher wear rates than females corroborating recent documentation of sex-specific life history tactics in ungulates. The data originate from two separated Danish roe deer populations exposed to contrasting habitats, but no difference in wear rate is found between populations. Thus, previous concern about the validity of age determination methods based on tooth wear may have been overstated. The findings demonstrate that objective measures of tooth wear can provide the basis for age determination in ungulate species that are otherwise difficult to age.     

Variations in adult body mass in roe deer: the effects of population density at birth and of habitat quality

Body mass is a key determinant of fitness components in many organisms, and adult mass varies considerably among individuals within populations. These variations have several causes, involve temporal and spatial factors, and are not yet well understood. We use long-term data from 20 roe deer cohorts (1977-96) in a 2600 ha study area (Chizé, western France) with two habitats contrasting in quality (rich oak forest in the North versus poor beech forest in the South) to analyse the effects of both cohort and habitat quality on adult mass (i.e. median body mass between 4 and 10 years of age) of roe deer (Capreolus capreolus). Cohort strongly influenced the adult body mass of roe deer in both sexes: males born in 1994 were 5.2 kg heavier when aged between 4 and 10 years old than males born in 1986, while females born in 1995 were 4.7 kg heavier between 4 and 10 years old than females born in 1982. For a given cohort, adult males were, on average, 0.9 kg heavier in the rich oak forest than in the poor beech forest. A similar trend occurred for adult females (0.5 kg heavier in the oak forest). The effects of cohort and habitat were additive and accounted for ca. 40% of the variation observed in the adult mass of roe deer at Chizé (males: 41.2%; females: 40.2%). Population density during the spring of the birth accounted for about 35% of cohort variation, whereas rainfall in May-June had no effect. Such delayed effects of density at birth on adult body mass probably affect population dynamics, and might constitute a mechanism by which delayed density-dependence occurs in ungulate populations.     

Effects of maternal age and condition on parturition and the perinatal period of Antarctic fur seals

The effect of maternal age and condition on the date of parturition and the duration of the perinatal period of Antarctic fur seals at Bird Island, South Georgia, were investigated over three consecutive breeding seasons. Females rear young during a four-month lactation period in a highly seasonal but predictable environment. Although females may first pup at three years of age, they did not attain full adult size until six years of age; older females (≥ 6 years) tended to be heavier, longer, and in better condition than younger females (3–5 years). Older females returned to breeding beaches earlier and could occupy the most suitable pupping sites, and gave birth when densities of animals on the beaches were low (i.e. more favourable for pup survival). Females that arrived earlier were able to remain ashore longer with their pups prior to departing on their first foraging trips but this was unrelated to either maternal age or condition. Younger females returned later in the pupping season, possibly as a result of late implantation due to smaller energy reserves than older and larger females. In 1990 all females arrived late, were in poorer condition, gave birth to lighter pups, and had shorter perinatal periods. This suggests that not only was implantation late but that females returned to an area of low food availability prior to parturition.     

How does environmental variation influence body mass,body size,and body condition? Roe deer as a case study

We tested the influence of population density and of drought intensity (measured as the Gaussen Index in spring and summer of the year of birth) on winter body mass, hind foot length, and body condition of roe deer fawns. Body mass decreased with increasing density and increased with increasing Gaussen Index in summer, in a similar way for both males and females. Hind foot length of males showed the same response. On the other hand, hind foot length of females decreased with increasing density only after dry summers, hence when environmental conditions were very harsh. Body condition was affected neither by density nor by drought intensity. Our results indicate that body mass and size are much better indicators of phenotypic quality than body condition in roe deer. The sex-specific responses of body size to environmental conditions could correspond to a differential allocation in favour of daughters by heavier than average roe deer mothers.     

Reproductive parameters of female Japanese macaques: Thirty years data from the arashiyama troops,Japan

Over a 30-year period from 1954 to 1983, 975 live births were recorded for Japanese macaque females at the Iwatayama Monkey Park, Arashiyama, Japan. Excluding unknown birth dates, primiparous mothers gave birth to 185 infants (182 cases with age of mother known) and multiparous mothers gave birth to 723 infants (603 cases with age of mother known). The peak month of birth was May with 52.3% of the total births occurring during the period. Multiparous females who had not given birth the previous year did so earlier than multiparous females who had given birth the previous year and also earlier than primiparous females. Among the females who had given birth the previous year, females whose infant had died gave birth earlier than females who had reared an infant the previous year. The offspring sex ratio (1:0.97) was not significantly different from 1:1, and revealed no consistent association with mother's age. Age-fecundity exhibited a humped curve. The annual birth rate was low at the age of 4 years but increased thereafter, ranging between 46.7% and 69.0%, at between 5 and 19 years of age, but again decreased for females between 20 and 25 years of age. Some old females displayed clear reproductive senescence. The infant mortality within the first year of age was quite low (10.3%) and the neonatal (less than 1 month old) mortality rate accounted for 49.0% of all infant deaths. There was no significant difference between the mortality rates of male and female infants. A female's rank-class had no apparent effect on the annual birth rate, infant mortality, and offspring sex ratio. These long-term data are compared with those from other primate populations.     

Density of Wild Prey Modulates Lynx Kill Rates on Free-Ranging Domestic Sheep

Understanding the factors shaping the dynamics of carnivore–livestock conflicts is vital to facilitate large carnivore conservation in multi-use landscapes. We investigated how the density of their main wild prey, roe deer Capreolus capreolus, modulates individual Eurasian lynx Lynx lynx kill rates on free-ranging domestic sheep Ovis aries across a range of sheep and roe deer densities. Lynx kill rates on free-ranging domestic sheep were collected in south-eastern Norway from 1995 to 2011 along a gradient of different livestock and wild prey densities using VHF and GPS telemetry. We used zero-inflated negative binomial (ZINB) models including lynx sex, sheep density and an index of roe deer density as explanatory variables to model observed kill rates on sheep, and ranked the models based on their AICc values. The model including the effects of lynx sex and sheep density in the zero-inflation model and the effect of lynx sex and roe deer density in the negative binomial part received most support. Irrespective of sheep density and sex, we found the lowest sheep kill rates in areas with high densities of roe deer. As roe deer density decreased, males killed sheep at higher rates, and this pattern held for both high and low sheep densities. Similarly, females killed sheep at higher rates in areas with high densities of sheep and low densities of roe deer. However, when sheep densities were low females rarely killed sheep irrespective of roe deer density. Our quantification of depredation rates can be the first step towards establishing fairer compensation systems based on more accurate and area specific estimation of losses. This study demonstrates how we can use ecological theory to predict where losses of sheep will be greatest, and can be used to identify areas where mitigation measures are most likely to be needed.     

Bigger teeth for longer life? Longevity and molar height in two roe deer populations

The role of tooth wear as a proximate cause of senescence in ruminants has recently been highlighted. There are two competing hypotheses to explain variation in tooth height and wear; the diet-quality hypothesis predicting increased wear in low-quality habitats, and the life-history hypothesis predicting molar height to be related to expected longevity. We compared tooth height and wear from roe deer of known age from two contrasting populations of roe deer (Capreolus capreolus) in France: Trois Fontaines (TF) with good habitat and shorter animal life expectancy and Chizé (CH) with poor habitat and longer animal life expectancy. There was no population difference in tooth wear, leading to rejection of the diet-quality hypothesis. However, despite their smaller body size, initial molar height for animals from CH was larger than for animals from TF. This provides the first evidence that variation in longevity between populations can lead to differences in molar height within a species.     

Population responses of roe deer to the recolonization of the French Vercors by wolves

In a context of changing carnivore populations worldwide, it is crucial to understand the consequences of these changes for prey populations. The recolonization by wolves of the French Vercors mountain range and the long-term monitoring (2001–2017) of roe deer in this area provided a unique opportunity to assess the effects of wolves on this prey. Roe deer was the main prey of wolves in the west Vercors mountain range during this recolonization. We compared roe deer abundance and fawn body mass in two contrasted areas of a wolf pack territory: a central area (core of the territory characterized by an intense use by wolves) and a peripheral area (used more occasionally). Roe deer population growth rates were lower in the central area between 2001 and 2006, resulting in a decline in roe deer abundance. Roe deer abundance substantially dropped in the two study areas after an extremely severe winter but the abundance of roe deer in the central area facing with wolves was slower to recover and remained at lower abundance levels for 6 years. Fawn body mass was consistently lower in the central area, varied similarly as roe deer abundance, and was not influenced by weather conditions or red deer population abundance. Altogether, the effects of wolves on roe deer in the central area occurred during a 10-year period following the establishment of wolves, through the interplay between wolf predation (before wolves started preying on red deer), harsh winter conditions and possibly naivety of prey to this recolonizing predator.     

Influence of Holocene environmental change and anthropogenic impact on the diversity and distribution of roe deer

Extant patterns of population structure and levels of diversity are a consequence of factors that vary in both space and time. Our objective in this study is to investigate a species that has responded to both natural and anthropogenic changes in ways that have shaped modern populations and provide insight into the key processes. The roe deer (Capreolus capreolus) is one of the two species of deer native to Britain. During the last glacial maximum (LGM), the British habitat was largely under ice and there was a land bridge to mainland Europe. As the Earth warmed during the early Holocene, the land bridge was lost. Subsequent hunting on the British mainland left the southern region extirpated of roe deer, whereas a refugial population remained in the north. Later reintroductions from Europe led to population expansion, especially in southern United Kingdom. Here, we combine data from ancient and modern DNA to track population dynamics and patterns of connectivity, and test hypotheses about the influence of natural and anthropogenic environmental change. We find that past expansion and divergence events coincided with a warming environment and the subsequent closure of the land bridge between Europe and the United Kingdom. We also find turnover in British roe deer haplotypes between the late-Holocene and modern day that have likely resulted from recent human disturbance activities such as habitat perturbation, overhunting and restocking.     

Fitness consequences of environmental conditions at different life stages in a long-lived vertebrate

The predictive adaptive response (PAR) hypothesis proposes that animals adjust their physiology and developmental trajectory during early life in anticipation of their future environments. Accordingly, when environmental conditions in early life match environmental conditions during adulthood, individual fitness should be greater. Here, we test this hypothesis in a long-lived mammal, the roe deer, using data from two contrasting populations, intensively monitored for more than 35 years. In the highly productive site, the fitness of female roe deer increased with the quality of environment during adulthood and, contrary to predictions of PAR, individuals born in good conditions always outperformed those born under poor conditions. In the resource-limited site, the fitness of female roe deer born in poor years was better than those born in good conditions in poor years when the animals were adult, but not in good years. Although consistent with predictions of PAR, we showed that this pattern is likely to be a consequence of increased viability selection during the juvenile stage for animals born in poor years. While PARs are often advanced in evolutionary medicine, our findings suggest that detailed biological processes should be investigated before drawing conclusions about the existence of this phenomenon.