Individual differences, density dependence and offspring birth traits in a population of red deer

Variation between individuals is an essential component of natural selection and evolutionary change, but it is only recently that the consequences of persistent differences between individuals on population dynamics have been considered. In particular, few authors have addressed whether interactions exist between individual quality and environmental variation. In part, this is due to the difficulties of collecting sufficient data, but also the challenge of defining individual quality. Using a long-established study population of red deer, Cervus elaphus, inhabiting the North Block of the Isle of Rum, and three quality measures, this paper investigates how differences in maternal quality affect variation in birth body mass and date, as population density varies, and how this differs depending on the sex of the offspring and the maternal quality measure used. Significant interactions between maternal quality, measured as a hind's total contribution to population growth, and population density are reported for birth mass, but only for male calves. Analyses using dominance or age at primiparity to define maternal quality showed no significant interactions with population density, highlighting the difficulties of defining a consistent measure of individual quality.     

Antler size in red deer: heritability and selection but no evolution

We present estimates of the selection on and the heritability of a male secondary sexual weapon in a wild population: antler size in red deer. Male red deer with large antlers had increased lifetime breeding success, both before and after correcting for body size, generating a standardized selection gradient of 0.44 (+/- 0.18 SE). Despite substantial age- and environment-related variation, antler size was also heritable (heritability of antler mass = 0.33 +/- 0.12). However the observed selection did not generate an evolutionary response in antler size over the study period of nearly 30 years, and there was no evidence of a positive genetic correlation between antler size and fitness nor of a positive association between breeding values for antler size and fitness. Our results are consistent with the hypothesis that a heritable trait under directional selection will not evolve if associations between the measured trait and fitness are determined by environmental covariances: In red deer males, for example, both antler size and success in the fights for mates may be heavily dependent on an individual's nutritional state.     

Antler growth rate in yearling Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>)

The aims of this study were to describe first antler growth rate and the factors affecting it. Growth rate reached a peak in week 14. Mean first antler growth rate during the whole growing period was 1.95 ± 0.05 cm/week. Lactation influenced antler growth rate as this was positively affected by total protein yield, while antlers grew at a lower rate the later they started to grow.     

Male fertility in natural populations of red deer is determined by sperm velocity and the proportion of normal spermatozoa

Male reproductive success is determined by the ability of males to gain sexual access to females and by their ability to fertilize ova. Among polygynous mammals, males differ markedly in their reproductive success, and a great deal of effort has been made to understand how selective forces have shaped traits that enhance male competitiveness both before and after copulation (i.e., sperm competition). However, the possibility that males also may differ in their fertility has been ignored under the assumption that male infertility is rare in natural populations because selection against it is likely to be strong. In the present study, we examined which semen traits correlate with male fertility in natural populations of Iberian red deer (Cervus elaphus hispanicus). We found no trade-offs between semen traits. Our analyses revealed strong associations between sperm production and sperm swimming velocity, sperm motility and proportion of morphologically normal spermatozoa, and sperm viability and acrosome integrity. These last two variables had the lowest coefficients of variation, suggesting that these traits have stabilized at high values and are unlikely to be related to fitness. In a fertility trial, our results show a large degree of variation in male fertility, and differences in fertility were determined mainly by sperm swimming velocity and by the proportion of morphologically normal sperm. We conclude that male fertility varies substantially in natural populations of Iberian red deer and that, when sperm numbers are equal, it is determined mainly by sperm swimming velocity and sperm morphology.     

Space use by red deer in a Mediterranean ecosystem as determined by radio-tracking

Space use, movements and activity rhythms of radio-tagged red deer were studied in a Mediterranean environment. The deer were found to be mainly crepuscular, nocturnal activity being higher than diurnal activity, which differs from studies of northern European populations of the species. Home ranges were smaller than most reported in the literature, that of the male being about two or three times larger than those of the females. Daily movements ranged from 3 to 4 km day⁻¹ according to the size of the home range. Vegetation cover was used more by the male and by all individuals during the summer, whilst open areas were used during the winter and spring, probably because of the high grass production in these seasons. The differences found with respect to other populations indicate that further studies in Mediterranean ecosystems are required to provide a basis of knowledge for management decisions.     

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.     

Step-size is determined by neck length in myosin V

The highly processive motor, myosin V, has an extremely long neck containing six calmodulin-binding IQ motifs that allows it to take multiple 36 nm steps corresponding to the pseudo-repeat of actin. To further investigate how myosin V moves processively on actin filaments, we altered the length of the neck by adding or deleting IQ motifs in myosin constructs lacking the globular tail domain. These myosin V IQ mutants were fluorescently labeled by exchange of a single Cy3-labeled calmodulin into the neck region of one head. We measured the step-size of these individual IQ mutants with nanometer precision and subsecond resolution using FIONA. The step-size was proportional to neck length for constructs containing 2, 4, 6, and 8 IQ motifs, providing strong support for the swinging lever-arm model of myosin motility. In addition, the kinetics of stepping provided additional support for the hand-over-hand model whereby the two heads alternately assume the leading position. Interestingly, the 8IQ myosin V mutant gave a broad distribution of step-sizes with multiple peaks, suggesting that this mutant has many choices of binding sites on an actin filament. These data demonstrate that the step-size of myosin V is affected by the length of its neck and is not solely determined by the pseudo-repeat of the actin filament.     

Sexual dimorphism and intercorhort variation in reindeer calf antler length is associated with density and weather

We analysed intercohort variability of live weight and antler length of 5,123 reindeer calves. We further assessed the influence of climate and density on the interannual variation in antler length, and discussed sex-specific resource allocation and response to climate variability. Antler length varied significantly among years and between sexes, with interaction between year and sex. Body weight and antler length were highly positively correlated, showed similar intercohort variability, and had a strong allometrical link, suggesting that antler length could be an equally reliable measure of calf condition as live weight. We found a relative measure of antler length (i.e. antler length corrected for the allometric effect of body mass) to be positively influenced by increasing density and May–June precipitation, and also decreasing May–June temperature. We attributed the effect of early summer weather to its influence on forage availability and quality as well as the level of parasitic insect harassment. Gender difference in both the allometric exponents and the interannual variability suggest that young males and females may have different tactics for relative resource allocation towards growth of antlers as compared to body mass. Because antlers are costly to produce, they may be an honest signal of individual quality for both sexes. However, we found gender-specific allometry, as female calves more than males appear to prioritize their antler growth over body mass, especially when resources are limited. Thus, our results suggest that environmental variation may influence the extent of sexual dimorphism in antler length. Electronic Supplementary Material Supplementary material is available for this article at     

Phylogeography,population genetics and conservation of the European red deer Cervus elaphus

• 1 During the Last Glacial Maximum, European red deer Cervus elaphus occurred in refugia in Iberia/southern France, Italy, the Balkans and the Carpathians. Most of Europe, including large parts of the east and north‐east, is now inhabited by red deer from the western lineage. The eastern lineage is largely confined to south‐eastern Europe; a third lineage comprises Sardo‐Corsican and Barbary red deer.
• 2 Sardo‐Corsican, Barbary and Mesola red deer are genetically unique units. They exhibit low levels of genetic diversity and deserve particular protection, since conservation strategies should target genetic information.
• 3 Hybridization between sika Cervus nippon and red deer occurs rarely, but may lead to extensive introgression, particularly in parts of the British Isles. Further expansion of both species may lead to increased hybridization in continental Europe.
• 4 Although hunting has an impact on red deer gene pools, the main threat today is habitat fragmentation in human‐dominated landscapes. The resulting increase in genetic drift and inbreeding reduces variability in isolated populations and may lead to inbreeding depression. To support vital meta‐populations, migration corridors should be established.

     

Antler growth and morphology in a feral sika deer (Cervus nippon) population in Killarney, Ireland

Antler growth and form was examined for a sample of 303 sika stags (Cervus nippon) culled over an eight-year period from a feral population in Killarney National Park, County Kerry, Ireland. Most antler measurements taken were highly correlated with each other and growth was complete after the sixth year. Few abnormalities were recorded and the degree of asymmetry in structure was less than previously reported for other sika populations. Variability in structure decreased with age and there was no effect of year of birth or cull on antler size for either adult or yearling stags. The growth pattern was similar to that of other populations except that the proportion of 'eight pointers' was low. An examination of antler damage was used to estimate fighting frequency across age classes. The results are discussed in relation to the genetic history of the herd and the use of antlers as weapons for intra-specific combat.     

Induction of early breeding in red deer (Cervus elaphus) by melatonin

Lactating (N = 12) and non-lactating (N = 6) red deer hinds and one stag at pasture were given concentrates (500 g/head) containing melatonin (5 mg/head) daily at 16:00 h from 18 June to 16 October. The stag shed the antler velvet and started rutting 5 weeks ahead of untreated stags, and hinds had their first oestrus and ovulation of the breeding season in mid-September, 5 weeks in advance of control lactating (N = 9) and non-lactating (N = 5) hinds. Treated hinds were allowed to mate with the treated stag and control hinds ran in an adjacent paddock with an untreated stag. All hinds became pregnant during the study with all but 2 melatonin-fed hinds (1 lactating, 1 non-lactating) and 1 control hind (lactating) conceiving at the first oestrus. Melatonin-treated hinds lost slightly more live weight than did controls from June to November, but their suckled calves grew throughout at a rate similar to those of control hinds.     

Disentangling demographic effects of red deer on chamois population dynamics

Investigating the impact of ecological factors on sex‐ and age‐specific vital rates is essential to understand animal population dynamics and detect the potential for interactions between sympatric species. We used block count data and autoregressive linear models to investigate variation in birth rate, kid survival, female survival, and male survival in a population of Alpine chamois Rupicapra rupicapra rupicapra monitored over 27 years within the Stelvio National Park, Central Italian Alps, as function of climatic variables, density dependence, and interspecific competition with red deer Cervus elaphus. We also used path analysis to assess the indirect effect of deer abundance on chamois growth rate mediated by each demographic parameter. Based on previous findings, we predicted that birth rate at [t] would negatively relate to red deer abundance at year [t − 1]; survival rates between [t] and [t + 1] would negatively relate to red deer abundance at year [t − 1] and to the interactive effect of winter precipitation at [t + 1] and chamois density at [t]. Our results showed that birth rate was positively related to spring–summer precipitation in the previous year, but this effect was hampered by increasing red deer abundance. Kid and female survival rates were negatively related to the combined effect of chamois abundance and winter precipitation. Male and female survival rates were negatively related to lagged red deer abundance. The path analysis supported a negative indirect effect of red deer abundance on chamois growth rate mediated by birth rate and female survival. Our results suggest that chamois population dynamics was largely explained by the synergistic effect of density dependence and winter harshness, as well as by interspecific competition with red deer, whose effects were seemingly stronger on the kid–female segment of the population.     

Interaction effects between weather and space use on harvesting effort and patterns in red deer

Most cervid populations in Europe and North America are managed through selective harvesting, often with age‐ and sex‐specific quotas, with a large influence on the population growth rate. Less well understood is how prevailing weather affects harvesting selectivity and off‐take indirectly through changes in individual animal and hunter behavior. The behavior and movement patterns of hunters and their prey are expected to be influenced by weather conditions. Furthermore, habitat characteristics like habitat openness are also known to affect movement patterns and harvesting vulnerability, but how much such processes affect harvest composition has not been quantified. We use harvest data from red deer (Cervus elaphus) to investigate how weather and habitat characteristics affect behavioral decisions of red deer and their hunters throughout the hunting season. More specifically, we look at how sex and age class, temperature, precipitation, moon phase, and day of week affect the probability of being harvested on farmland (open habitat), hunter effort, and the overall harvest numbers. Moon phase and day of week were the strongest predictors of hunter effort and harvest numbers, with higher effort during full moon and weekends, and higher numbers during full moon. In general, the effect of fall weather conditions and habitat characteristics on harvest effort and numbers varied through the season. Yearlings showed the highest variation in the probability of being harvested on farmland through the season, but there was no effect of sex. Our study is among the first to highlight that weather may affect harvesting patterns and off‐take indirectly through animal and hunter behavior, but the interaction effects of weather and space use on hunter behavior are complicated, and seem less important than hunter preference and quotas in determining hunter selection and harvest off‐take. The consideration of hunter behavior is therefore key when forming management rules for sustainable harvesting.     

Sex- and age-dependent effects of population density on life history traits of red deer Cervus elaphus in a temperate forest

We studied both the short‐ and long‐term effects of density on three life history traits of a red deer population inhabiting a temperate forest. Both male and female body mass increased when population density decreased, but male mass changed to a greater extent than female mass. Density did not influence female survival irrespective of age, however, survival of males was lower at high density for all age classes except the prime‐age class. Pregnancy rates of primiparous females increased markedly with decreasing density, whereas those of adult hinds were fairly constant and unrelated to density. For both sexes, of the studied life history traits we detected a long‐term effect of density at birth (cohort effect) only on body mass. These results suggest that density influences life history traits in the same way as factors of environmental variation such as climate. In this population we did not find any evidence for an influence of climatic conditions on life history traits of red deer. Both mild winters and the absence of summer droughts during the study period could account for such an absence of climatic effects. We interpreted our results to show that 1) as expected for a highly dimorphic and polygynous species such as red deer, male traits showed consistently higher sensitivity to variation in density than female traits, illustrating possible costs caused by sexual selection in males, 2) the female‐based Eberhardt's model according to which increasing density should sequentially affect juvenile survival, reproductive rates of primiparous females, reproductive rates of adults and lastly adult survival was only partly supported because we found that pregnancy rate of primiparous females rather than juvenile survival was the most sensitive trait to variation in density. We propose that including variation in male traits would improve the accuracy of models of population dynamics of large mammals, at least for highly dimorphic species. Because the population we studied was not fenced, we only measured apparent survival. We discuss how dispersal, in relation to the phenotypic quality of young deer, might be a potential regulating factor under such conditions.     

Patch occupancy, population density and dynamics in a fragmented red squirrel Sciurusvulgaris population

We studied population dynamics of red squirrels in a group of small forest fragments, that cover only 6.5% of the total study area (4664 ha) and where distances to the nearest source population were up to 2.2 km. We tested effects of patch size, quality and isolation and supplementary feeding on patch occupation during 1995–99. Larger patches and patches with supplementary feeding had a higher probability of being occupied. No patch <3.5 ha was ever occupied. No effects of isolation were found, suggesting that the forest habitat in the study area is not sufficiently fragmented to influence red squirrel distribution across patches. For medium sized patches (3.7–21 ha), that were occupied some years, there was an increase in patch occupation over the years, even though overall population size tended to decrease. These patches had a high turnover, especially of males. Patches in which the squirrel population went extinct were recolonized within a year. For patches that were at least some years occupied, squirrel density depended on patch quality only. No effects of patch size, isolation and winter temperature on population density were found. These data suggest that in our study area habitat fragmentation has no effect on local squirrel density and that the random sample hypothesis explains the distribution pattern across patches.     

Allometric growth, life-history invariants and population energetics

Population and community level processes must be at least partially determined by variation in the body sizes of constituent individuals, implying quantitative scaling relations can be extended to account for variation in those processes. Here we integrate allometric growth and life‐history invariant theories, and use this approach to develop theory describing the energetics of stationary populations. Our predictions approximate, with no free parameters, the scaling of production/biomass and assimilation/biomass ratios in mammalian populations and work partially for fish populations. This approach appears to be a promising direction and suggests the need for further development of the growth and life‐history models, and extensions of those theories.