Phenotypic and reproductive variation in female white-tailed deer: The role of harvest and environment

Animal density and stochastic environmental events affect physical development and recruitment of cervids. Biologists who manage cervid populations based on a density-dependent paradigm may need to also consider environmental effects. We analyzed 12- to 25-year time series of 3 harvested white-tailed deer populations in Mississippi to determine the relative influence of harvest and environmental factors on female reproduction and phenotypic quality. Using simple and multiple linear regression, we related body mass of 1.5-, 2.5-, and ≥3.5-year females, and 2.5- and ≥3.5-year percent lactation to variables representing deer harvest, growing season precipitation and temperatures, high-quality agronomic plantings, and flooding events. Response across populations varied greatly, with harvest variables explaining most, some, or no variation in body mass and percent lactation. Biologists should consider the potential influence of environmental factors on phenotypic and reproductive variation when making management decisions. © 2012 The Wildlife Society.     

Habitat quality influences relative antler size and hunters’ selectivity in roe deer

Understanding factors affecting antler size, and the extent to which harvesting is selective for these traits, is important in order to address management strategies aimed to minimize the risk of negative evolutionary consequences. In an Alpine study area, we compared the phenotypic quality and the antler size of 2,725 male roe deer hunted in two regions differing for winter harshness and habitat quality, and evaluated whether the selective behaviour of recreational hunters was influenced by phenotypic quality and antler size. Antler length and antler circumference relative to both body mass and jaw length were larger in the region with more favourable climate and habitat conditions, indicating that here roe deer were able to allocate more resources to antler growth. The analysis of the temporal trends of harvest bags suggested that hunters did not select roe deer for their body mass or size, but instead for antler size. This resulted also in a preference for sub-adult and adult age classes, while yearlings were culled reluctantly, especially in the region where antlers were smaller. Our results indicate that environmental heterogeneity may influence the relative investment in antler growth. In this way, it may interact with the hunters’ preferences increasing the risk that recreational hunting of roe deer, which is a widespread practice in many European countries, might result in alteration of male age structure and possibly in directional artificial selection.     

Contribution of dogs to white-tailed deer hunting success

Dogs (Canis familiaris) are used in hunting white-tailed deer (Odocoileus virginianus) in 10 North American jurisdictions. Although the practice is longstanding and controversial, the effects of dogs on the outcome of the hunt have rarely been studied. We evaluated the influence of dogs on recreational hunting of white-tailed deer based on long-term data from southeastern Ontario, Canada. Over 25 years, annual surveys of hunters were used to collect data on hunting effort and deer harvest from approximately 85 camps, roughly half of which had dogs. We investigated the relationship between harvest and 3 treatments (i.e., 0 dogs, 1 dog, and ≥2 dogs in camp), interactions with weather and deer density, and effects of neighboring camps. Dogs enhanced hunter success. We found no difference in deer encounter rates but, per unit effort, camps with ≥2 dogs harvested 0.013 (26%) more deer per hunter-day, missed 0.010 (23%) more deer per hunter-day, and wounded 0.002 (40%) more deer per hunter-day than camps without dogs. Conversely, camps without dogs saw, without shooting at, 0.033 (23%) more deer per hunter-day than camps with ≥2 dogs. These results are consistent with the idea that hunters with dogs are less selective. Hunters with dogs harvested more fawns per unit effort, but we found no difference in the harvest rate of older female deer. More precipitation, greater wind speed, lower temperatures and greater deer density improved harvest success but had no differential effect among dog treatments. Hunter success at camps with ≥2 dogs was less when neighboring camps also had ≥2 dogs. Because antlerless deer quotas are the principal means to control populations, increasing use of hunting dogs is unlikely to have substantial effects in managing overabundant deer. © 2012 The Wildlife Society.     

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     

Influence of weather and phenotypic characteristics on pregnancy rates of female roe deer in central Italy

In many ungulates, female fecundity is influenced by weather, population density and body condition. Based on a six-year survey of the reproductive tracts of adult female roe deer (Capreolus capreolus) harvested in the province of Pisa (Tuscany, central Italy), we evaluated the influence of weather, population density and individual characteristics on pregnancy rates. Eighty-five percent of females were pregnant, with a median litter size of two (range one—three). We found that pregnancy rate was positively correlated with summer rainfall and body mass of females, whereas early-winter conditions, spring rainfall, the age of females and density-dependent factors did not appear to influence pregnancy rate. These results reflect the particular seasonal variation in the abundance and quality of resources in Mediterranean habitats and show that heavier females (high-quality mothers) are more productive than lighter females.     

Evaluating how hunters see and react to telemetry collars on white-tailed deer

Fates of individuals outfitted with radiotransmitters commonly are used for estimating survival rates in populations of large animals that are hunted. Despite precautions, this practice may be subject to complex biases associated with hunter reaction to presence of radiotransmitters. To assess this potential bias we conducted an experiment using artificial deer (i.e., decoys) to measure hunters' abilities to see deer and determine if deer seen were wearing radiocollars. We used logistic regression to quantify probabilities that seeing deer and subsequently seeing radiocollars might be influenced by distance, percent visual obstruction, body orientation, hunter experience, and antler characteristics of deer. Additionally, we evaluated how experience and antler characteristics of deer might influence a hunter's decision to harvest a radiocollared deer. We found that 25.8% of the potentially observable collared deer (n = 663) were subsequently observed by hunters. Odds of observing deer and radiocollars increased 95% and 230%, respectively, for each additional log(yr) of hunting experience. Willingness to harvest radiocollared deer increased 89% for each additional log(yr) of hunting experience and 144% for large-antlered deer relative to antlerless deer. When hunting is an important source of mortality, analysts need to understand how potential biases associated with observing deer are associated with hunters' reactions to and subsequent decisions to harvest radiocollared animals. Our study suggested that presence of radiocollars may influence a deer's potential risk of being harvested and in turn bias telemetry-based estimates of survival, given that hunting mortality is the largest component of total mortality in hunted deer populations. Collar-based telemetry is used nearly universally by wildlife managers and researchers throughout North America and elsewhere to estimate and monitor the survival of big game populations that are managed through hunting. Our findings demonstrate that these estimates are likely subject to complex and systematic biases that managers should consider when evaluating future population-level effects of managed hunting. © 2011 The Wildlife Society     

Assessing the intensity of sexual selection on male body mass and antler length in roe deer Capreolus capreolus: is bigger better in a weakly dimorphic species?

Little is known about traits under sexual selection in territorial mammals with low sexual size dimorphism. We examined the potential for sexual selection on male body mass and antler length in the European roe deer Capreolus capreolus, a territorial ungulate in which males are less than 10% heavier than females. Independently, both body mass and antler length (irrespective of age) had a positive effect on male yearly breeding success. However, when corrected for body mass, antler length at a given mass only had a slight effect on male breeding success. This suggests that: (1) ‘bigger is better’ and (2) sexual selection is responsible for at least part of the observed variation in body mass and antler length in roe deer. High body mass and large antlers may be advantageous to males for two reasons: (1) they enhance fighting ability and dominance, so allowing males to defend better their territory and hence access mates, and (2) they attract females because they are honest signals of male phenotypic quality. This suggests that, even in weakly dimorphic ungulate species, sexual selection may lead to a marked influence of body mass on male breeding success, as long as body mass is also strongly selected in females.     

Influence of body mass and environmental conditions on winter mortality risk of a northern ungulate: Evidence for a late‐winter survival bottleneck

1. A relationship between winter weather and survival of northern ungulates has long been established, yet the possible roles of biological (e.g., nutritional status) and environmental (e.g., weather) conditions make it important to determine which potential limiting factors are most influential.
2. Our objective was to examine the potential effects of individual (body mass and age) and extrinsic (winter severity and snowmelt conditions) factors on the magnitude and timing of mortality for adult (>2.5 years old) female white‐tailed deer (Odocoileus virginianus [Zimmerman, 1780]) during February–May in the Upper Peninsula of Michigan, USA.
3. One hundred and fifty deer were captured and monitored during 2009–2015 in two areas with varying snowfall. February–May survival ranged from 0.24 to 0.89 (mean = 0.69) across years. Mortality risk increased 1.9% with each unit increase in cumulative winter severity index, decreased 8.2% with each cumulative snow‐free day, and decreased 4.3% with each kg increase in body mass. Age and weekly snow depth did not influence weekly deer survival. Predation, primarily from coyote (Canis latrans [Say, 1823]) and wolves (Canis lupus [L., 1758]), accounted for 78% of known‐cause mortalities.
4. Our results suggest that cumulative winter severity, and possibly to a lesser degree deer condition entering winter, impacted deer winter survival. However, the timing of spring snowmelt appeared to be the most influential factor determining late‐winter mortality of deer in our study. This supports the hypothesis that nutrition and energetic demands from weather conditions are both important to northern ungulate winter ecology. Under this model, a delay of several weeks in the timing of spring snowmelt could exert a large influence on deer survival, resulting in a survival bottleneck.

     

Habitat and weather influence body condition in white-tailed deer,Wisconsin, USA

In temperate and northern ecosystems where there are pronounced seasonal patterns in weather and available energy, there are corresponding patterns of body condition among white-tailed deer (Odocoileus virginianus). Body condition of white-tailed deer can affect survival and reproduction, which has large repercussions for state-level natural resource agencies that allocate hunting permits. In this study, we investigated how variation in winter weather, spring phenology, habitat composition, and browse quantity affected white-tailed deer body condition across a large spatial scale. Several body condition indicators (e.g., carcass mass, heart fat, antler size) were measured by hunters for 795 deer during September–December 2016–2018 in Wisconsin, USA. Winter severity in the previous year was an unreliable predictor of fall body condition of deer when winters were considered mild or moderate. The timing of spring green-up had a consistent effect on the body condition of all age and sex classes of deer. Earlier spring green-up resulted in heavier fawns and larger antlers among adult males. Region and spring green-up interacted to affect the heart fat of adult females. Earlier springs resulted in adult females in northern and central Wisconsin having a higher probability of heavy heart fat, whereas spring green-up had no effect on adult female heart fat in southern Wisconsin. Effects of habitat differed by age and sex class of deer, and by the body condition metric being evaluated, indicating that there are important physiological differences among age and sex classes of deer that are affected by the environment. Our study demonstrates that the hunting public can contribute large-scale, cost-effective, and quality data to deer monitoring and research projects. It is important that natural resource agencies be able to identify and recruit highly engaged members of the hunting public to ensure project success. The timing of spring green-up can have lasting effects on deer health that can be consistently observed the following fall, which is in contrast to the effects of winter severity that did not appear to persist when previous winters were mild or moderate. We encourage managers in northern or temperate regions to consider measures of spring green-up timing in conjunction with traditional winter severity when making deer population management decisions, such as antlerless tag allocation.     

Influence of Landscape Composition and Structure on Antler Size of White-Tailed Deer

Abstract: Cause for spatial variation in phenotypic quality of white-tailed deer (Odocoileus virginianus) populations is of great interest to wildlife managers. Relating phenotypic variation of populations to large-scale land-use patterns may provide insight into why populations exhibit spatial variation and elucidate how management can influence population phenotype. We used an information-theoretic approach to relate average antler size of 203 deer populations to composition and structure of the habitat occupied by those populations. We used interspersion, edge, and diversity indices to represent habitat structure and percentage of area in vegetation types to represent habitat composition. Landscape composition was a better predictor of deer population antler size than was landscape structure. Percentages of the management unit in agriculture, pasture, and pine forest were variables commonly found in the region-specific set of best models. Model-averaged estimates of agriculture and pasture parameters were always positive and estimates of pine forest parameters were always negative, which suggests that land-use types that promote growth of early successional herbaceous plants will positively influence antler size and, most likely, body growth and reproduction of white-tailed deer populations. Conversely, our findings suggest landscapes dominated by pine forests did not provide optimal amounts of quality forages for white-tailed deer. Pine forest effects should be mitigated using a combination of increased harvest to lower deer density and silvicultural practices like thinning, prescribed burning, and selective herbicide applications that stimulate growth of high-quality forages beneath the forest canopy to improve deer phenotypic quality.     

Seasonal changes in plasma leptin concentration related to antler cycle in Iberian red deer stags

The aim of this study is to describe the leptin cycle in male Iberian red deer (Cervus elaphus hispanicus) and relate it to antler and testosterone cycles. An additional aim is to assess the relationship between the plasma leptin concentration during antlers’ growth and their final size. Therefore, blood from 21 Iberian red deer males was sampled monthly to analyse leptin and testosterone. At the same time the deer were weighed and their body condition was assessed. The length of antlers was measured every 2 weeks and, after casting, their final length and perimeters were taken. Leptin showed a seasonal cycle, with a peak in June that decreased as testosterone increased. Low values were observed in autumn, winter and early spring. The relationship observed between leptin and body mass or body condition score was different in spring, when plasma testosterone concentration is low, than in autumn, when testosterone increases. Leptin peak amplitude was positively related to final antler size. In conclusion, the relationship between leptin and body mass and body condition score changes through the year, possibly due to the influence of androgens and photoperiod. There was a positive relationship between plasma concentration of leptin during antler growth and final antler length.     

Major-histocompatibility-complex-associated variation in secondary sexual traits of white-tailed deer (Odocoileus virginianus): evidence for good-genes advertisement

Abstract Good-genes hypotheses predict that development of secondary sexual characters can be an honest advertisement of heritable male quality. We explored this hypothesis using a cervid model (adult, male white-tailed deer, Odocoileus virginianus ) to determine whether antler development could provide an honest signal of a male's genetic quality and condition to adversaries. We compared antler, morphometric, hormonal, and parasitic data collected from hunter-harvested deer to characteristics of the Mhc-DRB (Odvi) , the most widely studied gene of the major histocom-patibility complex (MHC) in Artiodactyla. We detected associations between genetic characteristics at Odvi-DRB and antler development and body mass, suggesting that antler development and body mass may be associated with pathogen resistance in deer and thus may be an honest signal of genetic quality. We also detected associations between Odvi-DRB characteristics and serum testosterone during the breeding season, suggesting that certain MHC characteristics may help deer cope with stresses related to breeding activity. In addition, we observed a negative relationship between degree of antler development and overall abundance of abomasal helminths. Our observations provide support for the hypothesis that antler development in white-tailed deer is an honest signal of quality.     

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.     

Does fluctuating asymmetry of antlers in white-tailed deer (Odocoileus virginianus) follow patterns predicted for sexually selected traits?

Secondary sexual characters have been hypothesized to signal male quality and should demonstrate a negative relationship between the size of the trait and degree of fluctuating asymmetry because they are costly to produce. We collected morphometric and antler data from 439 white-tailed deer (Odocoileus virginianus) in Oklahoma, USA, in order to determine whether measures of antler asymmetry follow the patterns predicted for sexually selected characters. Relative fluctuating asymmetry was negatively related to antler size for all deer and within age groups up to five and a half years of age. We did not detect an association between asymmetry and antler size among deer that were six and a half years or older. When categorizing deer by antler size, we found that deer with small antlers (< or = 33rd percentile) had greater levels of relative asymmetry than deer with large antlers (< or = 67th percentile). The relative asymmetry of antlers was negatively related to age and was greatest in deer that were one and a half years old. Relative asymmetry was also negatively related to carcass mass, inside spread, skull length and body length. These data suggest that asymmetry in the antlers of white-tailed deer may be a reliable signal of quality and, as such, may be important in maintaining honesty in intrasexual advertisements during the breeding season.     

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 length of yearling red deer is determined by population density, weather and early life-history

In red deer, yearling antler length is a largely nutrition-mediated phenotypic character, and is therefore sometimes used as an indirect estimate of range quality and population condition. However, the parameters affecting yearling antler length have been little studied. We analyse the contributions of density, weather and maternal effects on yearling antler length of 581 individual stags born 1970-1996 on the Isle of Rum (Scotland). We show that antler length is a good measure of yearling condition: the probability of overwinter survival in yearlings that developed antlers was 3 times higher than for yearlings that did not develop antlers, and yearling antler length was correlated with the number of antler points the following year. Between years, variation in yearling antler length was best explained by variation in red deer density and June temperature at 12 months of age. Both of these variables were negatively correlated with antler length, and most likely this effect is due to changes in nutrient availability. Population density affects biomass availability for the individual, while low temperatures in early summer prolong the availability of high forage quality. At the individual level, antler length increased with birth weight and decreased with birth date, reflecting the persistent and pervasive influence of conditions in early life.     

Antler size provides an honest signal of male phenotypic quality in roe deer

Identifying factors shaping secondary sexual traits is essential in understanding how their variation may influence male fitness. Little information is available on the allocation of resources to antler growth in territorial ungulates with low sexual size dimorphism. We investigated phenotypic and environmental factors affecting both absolute and relative antler size of male roe deer in three contrasting populations in France and Sweden. In the three populations, we found marked age-specific variation in antler size, with an increase in both absolute and relative antler size between yearling and prime-age stages, followed by a decrease (senescence) for males older than 7 years. Antler size increased allometrically with body mass. This increase was particularly strong for senescent males, suggesting the evolution of two reproductive tactics: heavy old males invested particularly heavily in antler growth (potentially remaining competitive for territories), whereas light old males grew small antlers (potentially abandoning territory defense). Finally, environmental conditions had little effect on antler size: only population density negatively affected absolute antler size in one of the three populations. Antler size may therefore provide an honest signal of male phenotypic quality in roe deer. We discuss the implications of these results in terms of territory tenure and mating competition.     

Influence of Summer and Autumn Nutrition on Body Condition and Reproduction in Lactating Mule Deer

ABSTRACT Recent work suggests that availability and quality of forage in late summer and early autumn, a time when female ungulates face multiple energetic demands, is critical to reproduction in wild ungulates. Therefore, we examined direct links between nutritional quality of diets, body condition, and reproduction of lactating mule deer. Using captive mule deer, we tested the hypothesis that females consuming diets with lower digestible energy (DE; kJ/g) would have lower DE intake rates (DEI; MJ/day), have less body fat and muscle, have later estrus cycles, and have lower pregnancy and twinning rates. Deer fed lower DE diets had lower DEI during summer and autumn. In turn, deer with lower DEI, regardless of diet DE, had lower body mass, body fat, and muscle thickness. When nutritional quality of diets began to decline earlier in the summer, relationships between food quality, DEI, and body condition were stronger. Although DEI did not influence estrus date for deer that became pregnant before 21 December, deer with lower DEI had a lower probability of becoming pregnant and had a lower probability of producing twins. Measures of body condition in October (i.e., body mass, body fat, and muscle depth) predicted pregnancy and twinning rates in mule deer. Serum concentration of hormones leptin and Insulin Growth Factor 1 were not good predictors of body condition or reproduction. These findings suggest that managers concerned with productivity of mule deer populations should consider focusing on assessing and improving quality of forage available in summer and autumn.     

Population dynamics of Norwegian red deer: density-dependence and climatic variation.

We present a model on plant-deer climate interactions developed for improving our understanding of the temporal dynamics of deer abundance and, in particular, how intrinsic (density-dependent) and extrinsic (plants, climate) factors influence these dynamics. The model was tested statistically by analysing the dynamics of five Norwegian red deer populations between 1964 and 1993. Direct and delayed density-dependence significantly influenced the development of the populations: delayed density-dependence primarily operated through female density, whereas direct density-dependence acted through both female and male densities. Furthermore, population dynamics of Norwegian red deer were significantly affected by climate (as measured by the global weather phenomenon, the North Atlantic Oscillation: NAO). Warm, snowy winters (high NAO) were associated with decreased deer abundance, whereas the delayed (two-year) effect of warm, snowy winters had a positive effect on deer abundance. Our analyses are argued to have profound implications for the general understanding of climate change and terrestrial ecosystem functioning.     

Iodine Distribution in the Environment as a Limiting Factor for Roe Deer Antler Development

The iodine-containing hormones produced by the thyroid gland play a role in the complex neuro-hormonal regulation of antler development. The proper function of the thyroid depends on the adequate iodine supply of the organism, which is directly related to the iodine content of food and drinking water. The purpose of this study was to explore the connection between the iodine content of the water base, which has a strong correlation with the iodine concentration of environmental components available to animals, and the antler weight of roe deer (Capreolus capreolus) shot in Hungarian hunting areas. Using a general linear model, controlling for the collective effects of other environmental factors (deer population density, harvest rate, land use, and soil fertility information), the iodine content of the water base explained 51.4% of the total variance of antler weights. The results suggest that antler weights increase with increasing iodine concentration regardless of other factors; thus, the environmental iodine distribution can be a limiting factor suppressing roe deer performance assessed here as antler weight. Further experimental studies of controlled iodine uptake are needed to define the exact physiological iodine requirements of roe deer bucks.