Population-level response of coyotes to a pulsed resource event

From foraging theory, generalist predators should increase consumption of prey if prey availability increases. Pulsed resource events introduce a large influx of prey to predators that may exhibit a functional response of increased consumption rate on, or specialization to, this abundant food resource. We predicted that coyotes (Canis latrans) would respond functionally to numerical increases of neonate white-tailed deer (Odocoileus virginianus) during the pulsed resource event of parturition. We used howl surveys and deer camera surveys with occupancy modeling to estimate densities for coyotes, adult deer, and fawns, respectively, in Upper Peninsula of Michigan, USA, 2009–2011. We estimated biomass of adult and fawn deer consumed by coyotes during 2 periods [fawn limited mobility period (LMP) and social mobility period (SMP)] in May–August each year. Coyote densities were 0.32 and 0.37/km² for 2010–2011, respectively. Adult deer densities (3.7–3.9/km²) and fawn densities (0.6–1.3/km²) were similar across years. Overall, fawn hair occurrence in coyote scats was 2.3 times greater in LMP than SMP. Estimated consumption of fawns between periods (n = 157–880) by coyotes varied, suggesting a functional response, with increasing consumption of fawns relative to their availability. Coyotes, on average, consumed 2.2 times greater biomass of fawns than adults across years, and consumed 1.5 times greater fawn biomass, on average, during LMP than SMP. We suggest that consumption rates of coyotes is associated positively with increases in fawn density, and fawn consumption by coyotes follows predictions of optimal foraging theory during this pulsed resource event.     

Density-dependent responses of fawn cohort body mass in two contrasting roe deer populations

We investigated the influence of population density on juvenile body mass in two contrasting roe deer populations, in Sweden (Bogesund) and France (Chizé), in which density was monitored for ≥15 years. We investigated the effect of population density and climatic conditions on cohort performance. We predicted that: (1) body mass of growing fawns should be sensitive to environmental changes, showing marked between-year variation (i.e., cohort effects), (2) fawns in the less productive (weakly seasonal, weakly predictable summer weather) habitat of Chizé should show stronger density-dependent responses due to more severe food competition during summer than fawns in the more productive (markedly seasonal, moderately predictable summer weather) habitat of Bogesund, and (3) fawns at Bogesund should be heavier both in absolute terms and relative to their size than their conspecifics in Chizé due to a higher degree of fat accumulation in northern environments. In both study sites we found marked cohort variation and clear effects of density, with body mass varying by as much as 29% over years. While neither summer nor winter climate influenced fawn body mass at Bogesund, fawns tended to be lighter after summers with high temperatures at Chizé. In addition, fawns were heavier after acorn mast years experienced in utero at Bogesund. As expected, the strength of the density-dependent response of fawn body mass was greater at Chizé than at Bogesund. For a given density, male fawns were consistently heavier than females in both sites. Lastly, both sexes at Bogesund had higher absolute body mass and were larger for a given body size than in Chizé. Our results clearly demonstrate that absolute density is a poor predictor of roe deer performance and supports the view that habitat quality has an overwhelming importance for determining fawn body mass in roe deer populations.     

Survival of white-tailed deer fawns on Marine Corps Base Quantico

Some jurisdictions in the eastern United States have reduced harvest of white-tailed deer (Odocoileus virginianus) because of perceived declines in recruitment and population size over the last decade. Although the restoration of American black bears (Ursus americanus) and the colonization of coyotes (Canis latrans) have increased fawn predation in some areas, limited information exists on how temporally dynamic resources and weather influence fawn survival. Therefore, we evaluated fawn survival probability, cause specific mortality, and if factors such as oak (Quercus spp.) mast abundance, winter severity, precipitation, and landscape composition influenced mortality risk on Marine Corps Base Quantico in northern Virginia, USA, from 2008 to 2019. We tracked 248 fawns outfitted with very high frequency radio-collars and predation was the leading cause of mortality (n = 42; 45%). We estimated survival to 133 days and survival pooling all years (2008–2019) was 0.50 (95% CI = 0.42–0.60). Increased annual red oak (Quercus spp.) mast abundance from the previous fall reduced mortality hazard for fawns. The longevity of our study revealed a link between fawn survival and a specific maternal resource (red oak mast) only available during gestation. Our results highlight the importance of oak mast in eastern deciduous forests and, more broadly, overwinter maternal condition on white-tailed deer recruitment.     

White‐tailed deer exploit temporal refuge from multi‐predator and human risks on roads

Although most prey have multiple predator species, few studies have quantified how prey respond to the temporal niches of multiple predators which pose different levels of danger. For example, intraspecific variation in diel activity allows white‐tailed deer (Odocoileus virginianus) to reduce fawn activity overlap with coyotes (Canis latrans) but finding safe times of day may be more difficult for fawns in a multi‐predator context. We hypothesized that within a multi‐predator system, deer would allocate antipredation behavior optimally based on combined mortality risk from multiple sources, which would vary depending on fawn presence. We measured cause‐specific mortality of 777 adult (>1‐year‐old) and juvenile (1–4‐month‐old) deer and used 300 remote cameras to estimate the activity of deer, humans, and predators including American black bears (Ursus americanus), bobcats (Lynx rufus), coyotes, and wolves (Canis lupus). Predation and vehicle collisions accounted for 5.3 times greater mortality in juveniles (16% mortality from bears, coyotes, bobcats, wolves, and vehicles) compared with adults (3% mortality from coyotes, wolves, and vehicles). Deer nursery groups (i.e., ≥1 fawn present) were more diurnal than adult deer without fawns, causing fawns to have 24–38% less overlap with carnivores and 39% greater overlap with humans. Supporting our hypothesis, deer nursery groups appeared to optimize diel activity to minimize combined mortality risk. Temporal refuge for fawns was likely the result of carnivores avoiding humans, simplifying diel risk of five species into a trade‐off between diurnal humans and nocturnal carnivores. Functional redundancy among multiple predators with shared behaviors may partially explain why white‐tailed deer fawn predation rates are often similar among single‐ and multi‐predator systems.     

Forage quality's influence on mule deer fawns

In many mule deer (Odocoileus hemionus) populations, recruitment of fawns drives population dynamics. The quality of food available to females and their fawns in summer and autumn may play an important role in fawn recruitment. We examined direct links between digestible energy (DE) content of food and the DE intake of females on the nutrient concentration of milk and between the nursing behavior, DE intake, growth, and survival in captive mule deer fawns. We offered females and their fawns diets that simulated the natural decline in DE content of forage from mid-summer to late autumn in many western landscapes. Fawns fed a higher DE diet weighed 14% more at the onset of winter, had fewer unsuccessful nursing attempts, consumed milk with more protein and energy, and had higher survival than fawns fed a low DE diet. Differences between fawn performances among treatments were greatest when diet quality began decreasing earlier in the summer. Because our results indicate that summer and autumn nutrition is likely to influence fawn recruitment, wildlife biologists should include metrics for summer precipitation and late autumn fawn mass in population models, and land managers should focus on methods for improving the nutritional carrying capacity of summer and early autumn habitats. © 2011 The Wildlife Society.     

Predator densities and white-tailed deer fawn survival

Predation is the dominant source of mortality for white-tailed deer (Odocoileus virginianus) <6 months old throughout North America. Yet, few white-tailed deer fawn survival studies have occurred in areas with 4 predator species or have considered concurrent densities of deer and predator species. We monitored survival and cause-specific mortality from birth to 6 months for 100 neonatal fawns during 2013–2015 in the Upper Peninsula of Michigan, USA, while simultaneously estimating population densities of deer, American black bear (Ursus americanus), coyote (Canis latrans), bobcat (Lynx rufus), and gray wolf (Canis lupus). We estimated fawn predation risk in response to sex, birth mass, and date of birth. Six-month fawn survival pooled among years was 36%, and fawn mortality risk was not related to birth mass, date of birth, or sex. Estimated mean annual deer and predator densities were 334 fawns/100 km², 25.9 black bear/100 km², 23.8 coyotes/100 km², 3.8 bobcat/100 km², and 2.8 wolves/100 km². Despite lower estimated per-individual kill rates, coyotes and black bears were the leading sources of fawn mortality because they had greater densities relative to bobcats and wolves. Our results indicate that the presence of more predator species in a system is not entirely additive in its effect on fawn survival. © The Wildlife Society, 2019     

Culling Versus Density Effects in Management of a Deer Population

Abstract: Wildlife managers often manipulate hunting regulations to control deer populations. However, few empirical studies have examined the level of hunting effort (hunter-days) required to limit population growth and demographic effects through harvesting of females. Moreover, the relative importance of density effects on population growth has not been quantified. We reconstructed a sika deer [Cervus nippon] population over a period of 12 years (1990–2001) using age- and sex-specific harvest data. Using cohort analysis, we analyzed population dynamics, focusing on 1) the relationship between hunting effort and hunting-induced mortality rate, 2) relative contributions of hunting mortality and recruitment of yearlings to annual changes in population growth rate, and 3) annual variation in recruitment rate. Population size increased until 1998 and declined thereafter. The population growth rate changed more in response to annual changes in recruitment rate than hunting mortality rate. Temporal variation in recruitment rate was not controlled by birth rate alone; direct density dependence, intensities of hunting mortality for fawns, and for females (≥2 yr of age), which accounted for the fawn survival rate, were required as factors to explain temporal variation. Density effects on the recruitment rate were not strong enough to regulate the population within the study period; high hunting mortality, with intensive female harvesting, was necessary to prevent population growth. Hunting effort was a good predictor of the hunting mortality rate, and female harvest had a negative effect on the recruitment rate through fawn survival. We suggest that >3,500 hunter-days and prioritization of female harvesting are required to prevent increases in this deer population.     

White-Tailed Deer Population Dynamics Following Louisiana Black Bear Recovery

Changing predator communities have been implicated in reduced survival of white-tailed deer (Odocoileus virginianus) fawns. Few studies, however, have used field-based age-specific estimates for survival and fecundity to assess the relative importance of low fawn survival on population growth and harvest potential. We studied white-tailed deer population dynamics on Tensas River National Wildlife Refuge (TRNWR) in Louisiana, USA, where the predator community included bobcats (Lynx rufus), coyotes (Canis latrans), and a restored population of Louisiana black bear (Ursus americanus luteolus). During 2013–2015, we radio-collared and monitored 70 adult (≥2.5 yrs) and 21 yearling (1.5-yr-old) female deer. Annual survival averaged 0.815 (95% CI = 0.734–0.904) for adults and 0.857 (95% CI = 0.720–1.00) for yearlings. We combined these estimates with concurrently collected fawn survival estimates (0.27; 95% CI = 0.185–0.398) to model population trajectories and elasticities. We used estimates of nonhunting survival (annual survival estimated excluding harvest mortality) to project population growth (λ) relative to 4 levels of harvest (0, 10%, 20%, 30%). Finally, we investigated effects of reduced fawn survival on population growth under current management and with elimination of female harvest. Despite substantial fawn predation, the deer population on TRNWR was increasing (λ = 1.06) and could sustain additional female harvest; however, the population was expected to decline at 20% (λ = 0.98) and 30% (λ = 0.94) female harvest. With no female harvest, the population was projected to increase with observed (λ = 1.15) and reduced fawn survival (λ = 1.02), but the population could not sustain current female harvest (10%) if fawn survival declined (λ = 0.90). For all scenarios, adult female survival was the most elastic parameter. Given the importance of adult female survival, the relative predictability in response of adult survival to harvest management, and the difficulty in altering fawn survival, reducing female harvest is likely the most efficient approach to compensate for low fawn survival. On highly productive sites such as ours, reduction, but not necessarily elimination, of harvest can mitigate effects of low fawn survival on population growth. © 2020 The Wildlife Society.     

Increased brown bear predation on sika deer fawns following a deer population irruption in eastern Hokkaido, Japan

By the 1970s, brown bears (Ursus arctos) in Hokkaido, northern Japan, were opportunistic omnivores that mainly depended on plant materials. Because the sika deer (Cervus nippon) population irrupted in eastern Hokkaido in the 1990s, we expected that brown bears might prey on sika deer fawns. First, we developed a simple and cost-effective method of monitoring possible bear predation on deer fawns by analyzing the widths of deer hairs remained in bear scats. Based on hair thickness standards, we distinguished the brown bear consumption of deer fawns from adults by analyzing bear scats (n?=?108) collected during the deer birthing season (late May?Clate July) in 1999?C2008. To evaluate the importance of fawns to bears, we compared the occurrence of fawn and adult deer hairs in bear scats among three periods (I, 1999?C2000; II, 2003?C2005; III, 2006?C2008) in eastern Hokkaido. The occurrence of fawn hairs in bear scats increased from 12.5 to 27.3?% in volume and from 6.3 to 33.6?% in frequency from period I to period III, whereas adult hairs in scats decreased from 42.8 to 26.1?% in volume and from 34.4 to 22.7?% in frequency during the same time. These data suggest that bears increasingly preyed on deer fawns after the deer population irruption and decreasingly used adult carcasses because of the enforcement of deer carcass treatment by the Hokkaido government.     

MULE DEER SURVIVAL AMONG ADJACENT POPULATIONS IN SOUTHWEST IDAHO

Abstract: We investigated survival and cause-specific mortality of mule deer (Odocoileus hemionus) on 3 distinct winter ranges in southwest Idaho from 1992 to 1997 to identify demographic variation and potential limiting factors based on a sample of 447 radiocollared deer. During winters 1995–1996 and 1996–1997, we modeled overwinter fawn mortality based on early winter mass, sex, activity, and habitat use variables. Annual survival rates of adult mule deer varied among the 3 adjacent study areas (χ₂² = 10.93, P = 0.004). Overwinter deer survival also varied among study areas (χ₂² = 8.00, P = 0.018), and the study area X year, study area X sex, and study area X age interactions were all significant (P ≤ 0.018). Overwinter survival differences among the study areas were not consistent over time or among sexes and ages of deer. Winter malnutrition was the main cause of mortality for both adults and fawns during the severe winter of 1992–1993, when overall survival was low. Excluding harvest, predation was the major proximate cause of deer mortality during 1993–97 when overall survival was higher. The probability of winter fawn mortality increased with lower mass (χ₂¹ = 7.38, P = 0.007), being male (χ₂¹ = 5.61, P = 0.018), smaller group sizes (χ₂¹ = 3.62, P = 0.057), and using steeper slopes (χ₂¹ = 3.05, P = 0.081). Smaller group sizes and use of steep slopes corresponded to conditions where predators were more successful. Our findings suggest that coyote (Canis latrans) predation was largely compensatory whereas mountain lion (Puma concolor) predation was apparently independent of animal condition and dependent more on deer habitat use. Early winter fawn mass was a better predictor of overwinter fawn survival than a suite of winter resource use variables, lending further support for use of fawn mass to predict winters where fawn mortality may be high. No single population in this study could be used to make reliable inferences regarding deer survival in the other populations. Survival rate measurements should be used cautiously to make inferences in populations where survival has not been directly measured.     

Sarcocystis hemionilatrantis (Sp. N.) life cycle in mule deer and coyotes.

Fifteen coyotes (Canis latrans) shed sporulated sporocysts in their feces after eating freshly ground skeletal muscles from a mule deer (Odocoileus hemionus hemionus) infected with microscopic-sized cysts of Sarcocystis. Sporocysts were shed intermittently from 12 to 36 days after ingestion of the infected meat. Sporocyst size averaged 14.4 X 9.3 mum. Eleven mule deer fawns orally inoculated with these sporocysts became infected and 9 of 11 died between post-inoculation days (PID) 27 and 63. Clinical signs of anorexia, weight loss, pyrexia and weakness were evident prior to death. A calf (Bos taurus) and two lambs (Ovis aries) orally inoculated with these sporocysts did not become infected and remained healthy throughout the experiments. Similarly, uninoculated control animals consisting of three mule deer fawns, two lambs and one calf remained healthy during the experiment. Preliminary histologic examinations conducted on selected tissues from all animals revealed microscopic-sized schizogonous stages in macrophages, between muscle fibers and near blood vessels in the esophagus, heart, biceps femoris, semi-membranosus, diaphragm and tongue from seven of eight fawns which died between PID 27 and 39. Developing or mature muscle cysts were not found in fawn tissue until PID 60. Sarcocysts were found in the three infected fawns examined after this time. Muscle cysts or earlier schizont stages were not found in tissues from the inoculated or uninoculated calves and lambs. A single muscle cyst was found in one control fawn; the other two control fawns were negative for both muscle cysts and other schizogonous stages. These results established that the life cycle of this species of Sarcocystis can be completed with coyotes as the definitive host and mule deer as the intermediate host. Based on the demonstrated host specificity and earlier findings, the name Sarcocystis hemionilatrantis is proposed for this parasite of mule deer and coyotes.     

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.     

Spatial variation in springtime food resources influences the winter body mass of roe deer fawns

It is well established that the dynamics of mammalian populations vary in time, in relation to density and weather, and often in interaction with phenotypic differences (sex, age and social status). Habitat quality has recently been identified as another significant source of individual variability in vital rates of deer, including roe deer where spatial variations in fawn body mass were found to be only about a tenth of temporal variations. The approach used was to classify the habitat into blocks a priori, and to analyse variation in animal performance among the predefined areas. In a fine-grained approach, here we use data collected over 24 years on 1,235 roe deer fawns captured at known locations and the plant species composition sampled in 2001 at 578 sites in the Chizé forest to determine the spatial structure at a fine scale of both vegetation and winter body mass of fawns, and then to determine links between the two. Space and time played a nearly equal role in determining fawn body masses of both sexes, each accounting for about 20% of variance and without any interaction between them. The spatial distribution of fawn body mass was perennial over the 24 years considered and predicted values showed a 2 kg range according to location in the reserve, which is much greater than suggested in previous work and is enough to have strong effects on fawn survival. The spatial distribution and the range of predicted body masses were closely similar in males and females. The result of this study is therefore consistent with the view that the life history traits of roe deer are only weakly influenced by sexual selection. The occurrence of three plant species that are known to be important food items in spring/summer roe deer diets, hornbeam (Carpinus betulus), bluebell (Hyacinthoides sp.) and Star of Bethlehem (Ornithogalum sp.) was positively related to winter fawn body mass. The occurrence of species known to be avoided in spring/summer roe deer diets [e.g. butcher's broom (Ruscus aculeatus) and beech (Fagus sylvatica)], was negatively related to fawn body mass. We conclude that the spatial variation in the body mass of fawns in winter in this forest is as important as the temporal variation, and that the distribution of plant species that are actively selected during spring and summer is an important determinant of spatial variation in winter fawn body mass. The availability of these plants is therefore likely to be a key factor in the dynamics of roe deer populations.     

饲养林麝的种群特征和幼麝的存活研究

对饲养在上海崇明岛东平林场的林麝种群特征以及1998～2000 年间该种群所出生的74 只幼麝存活情况进行分析研究。结果表明: 该林麝种群中能繁殖的个体占51.1 % , 种群数量逐渐增长。种群存活率曲线呈C 型,说明林麝的死亡主要发生在幼麝时期。从不同年龄段性比变化来看, 初生幼麝的性比接近1∶1 , 由于雌幼麝比雄幼麝具有更高的存活率( P < 0.05) , 进入亚成体以后雄性比例逐渐下降, 显示幼麝时期存在偏雄性死亡。早出生幼麝的存活率明显高于晚出生幼麝( P < 0.05) 。引起幼麝死亡的主要原因是疾病(45. 9 %) 。     

Vole fluctuations,red fox responses,predation on fawns,and roe deer dynamics in a temperate latitude

The alternative prey hypothesis predicts that predators respond both functionally and numerically (with a time lag) to fluctuations in the main prey abundance, which affects the survival of alternative prey. This pattern was found in northern Europe in the community formed by voles (Microtidae), red foxes (Vulpes vulpes) and roe deer (Capreolus capreolus). We studied the same predator—prey community in a temperate latitude where, according to the predation hypothesis, only the functional response of predators to changes in main prey availability should occur. In the years 1997–2007, in western Poland, we estimated the index of common vole (Microtus arvalis) abundance (burrow counts), the density of foxes (spotlight counts), the young production in foxes (young/adult ratio), the index of fox predation on fawns (prey remains near dens) as well as the reproduction index (fawn/female ratio) and density of roe deer (total counts). The vole abundance fluctuated considerably, the young production in foxes did not correlate with the main prey availability, but the density of foxes showed direct numerical response. The index of fox predation on fawns decreased with the vole abundance and negatively affected the fawn/female ratio in roe deer. Thus, the relationships between voles and foxes were not fully consistent with the predation hypothesis. The direct numerical response of foxes should tend to stabilize this predator—prey community. It is suggested, however, that responses showed by vole-eating predators in temperate latitudes may sometimes affect their alternative prey, including animals with unfavourable conservation status.     

Factors Influencing Survival Rates of Pronghorn Fawns in Idaho

Pronghorn (Antilocapra americana) occur throughout western North America. In Idaho, USA, following intensive hunting to reduce crop depredations in the late 1980s, pronghorn populations have not rebounded to desired levels. Because neonatal survival in ungulates is one factor limiting population growth, we evaluated cause-specific mortality and the influence of intrinsic and extrinsic factors on survival rates of 217 radio-collared pronghorn fawns across 3 study areas in Idaho during 2015–2016. For intrinsic variables, we determined the sex and body mass index (BMI) for each fawn. For extrinsic variables, we determined the abundance of predators and alternate prey, estimated the normalized difference vegetation index (NDVI) for 1 month pre- and post-parturition, and measured fecal nitrogen and diaminopimelic acid (DAPA). We considered NDVI as a measure of plant productivity, and fecal nitrogen and DAPA as possible proxies of diet quality. We predicted NDVI, fecal nitrogen, and DAPA would be positively related to the nutritional status of females and positively related to fawn survival. We used Program MARK with known fate models to estimate semi-monthly survival rates of pronghorn fawns for the first 4 months post-parturition. During both years, the leading cause of fawn mortality was coyote (Canis latrans) predation (58%), followed by unknown causes of mortality (18%), unknown predation (12%), predation by bobcats (Lynx rufus; 6%), predation by golden eagles (Aquila chrysaetos; 3%), and other (3%). Mean fawn survival for the 4 months post-parturition across years and study sites was 0.42 ± 0.04 (SE; range = 28–62%). The top survival model included BMI, lagomorph abundance, and DAPA and had a model weight of 83.3%. All 3 variables were positively related to pronghorn fawn survival. Because females with increased nutrition generally have heavier fawns, BMI was likely correlated to diet quality, which was supported by the positive relationship between DAPA and fawn survival. We hypothesize that high lagomorph abundance created an alternate prey base to buffer coyotes from preying on pronghorn neonates. We found no influence of measures of NDVI (pre- and post-parturition), fecal nitrogen, or predator abundance on fawn survival. Management actions providing high-quality forage for pronghorn are likely to contribute to production of heavier fawns having the highest chance of survival. © 2020 The Wildlife Society.     

Cyclic voles, prey switching in red fox, and roe deer dynamics – a test of the alternative prey hypothesis

Medium-sized predators sometimes switch to alternative prey species as their main prey declines. Our objective of this study was to test the alternative prey hypothesis for a medium sized predator (red fox, Vulpes vulpes ), a small cyclically fluctuating main prey (microtine voles) and larger alternative prey (roe deer fawns, Capreolus capreolus ). We used long-term time series (28 years) on voles, red fox and roe deer from the Grimsö Wildlife Research Area (59°40'N, 15°25'E) in south-central Sweden to investigate interspecific relationships in the annual fluctuations in numbers of the studied species. Annual variation in number of roe deer fawns in autumn was significantly and positively related to vole density and significantly and negatively related to the number of fox litters in the previous year. In years of high vole density, predation on roe deer fawns was small, but in years of low vole density predation was more severe. The time lag between number of fox litters and predation on fawns was due to the time lag in functional response of red fox in relation to voles. This study demonstrates for the first time that the alternative prey hypothesis is applicable to the system red fox, voles and roe deer fawns.     

Spatial patterns and density dependence in the dynamics of a roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) population in central Italy

The demography of roe deer living in a mountain area of central Italy was studied from 1995 to 1999 with 104 radio-tagged animals, including fawns and adults of both sexes. From spring surveys we estimated population density by mark-resighting (average: 53.8dž.8 individuals km^-2) and found an average fawn/doe ratio of 0.75ǂ.4. The fawn/doe ratio was negatively correlated to density suggesting density-dependent regulation in this population. Using culled and net-trapped individuals we evaluated the dressed body weights of adult males (23.1ǃ.0 kg) and females (22.0ǃ.0 kg), which indicated a low level of sexual dimorphism. The potential litter size (1.44ǂ.1 embryos) depended on female body weight and a threshold of 20.9ǃ.4 kg separated adult females carrying one or two embryos. Both fawn (0.38ǂ.07) and adult survival (0.90ǂ.07) were evaluated from radiotagged individuals and no gender effect was observed in either age class. During the study period we recorded a population decline in one part of the study area and an increasing fawn mortality, which was attributed to the spreading of an enteropathogenic desease. The study revealed an unexpected spatial structure in population dynamics at a scale of few square kilometres. In the two studied subareas, which are very close and ecologically similar, we documented significant differences in several demographic parameters: females in the subarea with the highest deer density produced smaller litters and allocated their reproductive effort preferentially to males, which is consistent with the hypothesis that local resource competition determines sex allocation in roe deer. The importance of spatial variability with respect to roe deer demography was overlooked in previous studies and our results raise new interesting research questions relative to the study of population equilibria which are also relevant for the management of this important game species.     

Winter fawn survival in black-tailed deer populations affected by hair loss syndrome

Overwinter fawn mortality associated with hair loss syndrome (HLS) is anecdotally thought to be important in declines of Columbian black-tailed deer (Odocoileus hemionus columbianus) populations in Washington and Oregon (USA). We determined prevalence of HLS in black-tailed deer, September and April fawn:doe ratios, and minimum overwinter survival rates of fawns for selected game management units (GMUs) in western Washington from 1999 to 2001. Prevalence of HLS ranged from 6% to 74% in fawns and 4% to 33% in does. Minimum fawn survival ranged from 0.56 to 0.83 and was unrelated to prevalence of HLS in either does (r=0.005, P=0.991) or fawns (r=-0.215, P=0.608). The prevalence of HLS in either does or fawns was also unrelated to either fall fawn:doe ratios (HLS does: r=-0.132, P=0.779; HLS fawns: r=0.130, P=0.760) or spring fawn:doe ratios (HLS does: r=-0.173, P=0.711; HLS fawns: r=-0.020, P=0.963). However, the prevalence of HLS in does and fawns was strongly related (r=0.942, P=0.002), and GMUs with high prevalence of HLS had lower deer population densities (fawns: r=-0.752, P=0.031; does: r=-0.813, P=0.026). Increased overwinter mortality of fawns because of HLS was not supported by our data. Decreased production of fawns, increased summer mortality of fawns, or both were seen in six of eight study GMU-year combinations. Observed rates of productivity and minimum fawn survival were inadequate to maintain population size in five of eight study GMU-year combinations, assuming an annual doe survival rate of 0.75. The influence of deer condition and population health on adult survival, fawn production, preweaning fawn survival, parasitism, and prevalence of HLS in both fawns and adults need to be clarified to identify what factors are limiting black-tailed deer productivity.     

Fight or flight? Antipredator behavior and the escalation of coyote encounters with deer

It is well known that prey of different size and morphology often use different antipredator strategies. The prevailing notion is that this occurs because size, morphology and weaponry determine the relative effectiveness of alternative strategies, and nowhere is this assumption more entrenched than in our view of the basic decision to stay, fight or flee. Here, we use observations of coyote (Canis latrans) packs hunting deer in winter to show that two ungulates of similar size and morphology, white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus), use different antipredator strategies when encountered or attacked. Mule deer typically responded by holding their ground and aggressively defending conspecifics, and were at high risk of being attacked and killed if they fled or were undefended. White-tails always fled when pursued or attacked by coyotes. Coyotes pursued fewer white-tails than mule deer they encountered regardless of prey response. Once pursued or attacked, white-tails faced a risk of attack and capture, respectively, that was intermediate between the high and low risk mule deer groups. The overall risk of capture per encounter for white-tails was similar to that facing mule deer that confronted coyotes, which was much lower than risk facing mule deer that fled and were undefended. Contextual variables such as the opportunity to improve one's position by joining another group, moving to rugged terrain, or the presence of companions that are willing to provide defense may explain why a mixed strategy is maintained in mule deer, despite the apparently detrimental effects of flight. These examples illustrate the value of including prey behavior in models of hunting success in so far as prey defenses may not be coupled with differences in size and morphology.