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.     

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.     

Predation processes: behavioural interactions between red fox and roe deer during the fawning season

Predation by red fox (Vulpes vulpes) is the most important mortality cause for neonatal roe deer (Capreolus capreolus) in Scandinavia. With the objective of investigating how the fox finds fawns and how antipredatory behaviour of roe deer females influences choice of hunting method, I analysed observations of interactions between red fox and roe deer females. The observations were collected over 14 years in a mixed forest/agricultural landscape in Sweden. Of 49 fox–doe encounters, the doe attacked the fox in 59%. In 90% of these attacks the fox was successfully deterred. In two observations a doe saved a fawn attacked by a fox. Two hunting methods used by the fox were discerned. In 28 cases foxes searched the ground, and in 18 cases they surveyed open areas, often from a forest edge. The latter behaviour seemed more directed at fawns and was seen leading to a capture attempt. Searching seemed less efficient and also difficult to conduct due to the aggressiveness of does. A surveying sit-and-wait type of hunting method thus appeared as the most successful. The possibility to use this method could explain why roe deer fawns are more vulnerable to fox predation in open habitats.     

Antibodies to granulocytic Ehrlichia in moose, red deer, and roe deer in Norway

Serum samples from 104 moose (Alces alces), 124 red deer (Cervus elaphus) and 114 roe deer (Capreolus capreolus), collected from different counties in southern Norway from 1994 to 2000, were analysed by an indirect immunofluorescent antibody staining method for antibodies to Ehrlichia equi. The overall seroprevalences for granulocytic Ehrlichia spp. in moose, red deer, and roe deer from Ixodes ricinus infested counties were 43%, 55%, and 96%, respectively. Antibody prevalence was significantly higher in roe deer than in moose and red deer (P < 0.001). Mean antibody titers (log10 +/- SD) to E. equi in sera from moose, red deer, and roe deer were 1:1,497 (3.17 +/- 0.646), 1:234 (2.37 +/- 0.424) and 1:676 (2.83 +/- 0.404), respectively. The present work indicates that all these wild ruminant species are exposed to granulocytic Ehrlichia in Norway.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Seasonal variation of activity patterns in roe deer in a temperate forested area

We investigated the activity patterns of a European roe deer (Capreolus capreolus) population living in a forested Apennine area in central Italy, in order to shed light on the environmental and biological factors that were expected to account for the observed activity patterns on daily and yearly bases. Daily and seasonal activity patterns of 31 radio-collared roe deer were assessed through sessions of radio tracking for a total period of 18 consecutive months. Roe deer showed bimodal activity patterns throughout the year, with the two highest peaks of activity recorded at dawn and dusk. Activity patterns of males and females differed during the territorial period (from early spring to late summer), whereas they did not during the nonterritorial period. Most likely, behavioral thermoregulation can be held responsible for variation of daily activity patterns in different seasons. In winter, for instance, activity during the dawn period was significantly higher than in other seasons and daylight activity was significantly higher than at night. Nocturnal activity was highest in summer and lowest in winter. During the hunting season, moreover, roe deer showed lower activity levels than during the rest of the year. The prediction that roe deer would show lower activity levels during full moon nights, when the predation risk was assumed to be higher, was not confirmed by our data. Activity rhythms in roe deer were thus subjected to both endogenous and environmental factors, the latter working as exogenous synchronization cues. Accordingly, in changing environmental and ecological conditions, a circadian cycle of activity could be seen as the result of complex interactions among daily behavioral rhythm, digestive physiology, and external modifying factors.     

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.     

Season effect on genitalia and epididymal sperm from Iberian red deer, roe deer and Cantabrian chamois

Seasonality deeply affects the physiology and behavior of many species, and must be taken into account when biological resource banks (BRBs) are established. We have studied the effect of seasonality on many reproductive parameters of free-ranging Iberian red deer, roe deer and Cantabrian chamois, living in Spain. Testicles from hunted animals were collected and sent to our laboratory at different times during the year. We recorded the weight and volume of testis, the weight of the epididymis and its separate parts (caput, corpus, and cauda), the weight of the sperm sample collected from the cauda epididymis, and several sperm parameters (sperm concentration, spermatozoa recovered, motility, HOS test reactivity, acrosomal status, and viability). We studied the data according to several periods, defined accordingly to each species. For red deer, we defined rut (mid-September to mid-October), post-rut (mid-October to mid-December), and non-breeding season (February). For roe deer, they were pre-rut (June), rut (July), post-rut (first fortnight of August), and non-breeding season (September). For chamois: non-breeding season (June to mid-September) and breeding season (October-November). The rut/breeding season yielded significantly higher numbers for almost all parameters. However, in the case of red deer, sperm quality was higher in the post-rut. For roe deer, testicular weight was similar in the pre-rut and in the rut, and sperm quality did not differ significantly between these two periods, although we noticed higher values in the rut. In the case of chamois, sperm quality did not differ significantly from the breeding season, but data distribution suggested that in the non-breeding season there are less males with sperm of good quality. On the whole, we find these results of interest for BRB planning. The best season to collect sperm in this species would be the breeding season. However, post-rut in red deer, pre-rut in roe deer, and non-breeding season in chamois could be used too, because of the acceptable sperm quality, despite the lower quantity salvaged. More in-depth research needs to be carried out on the quality of sperm salvaged at different times of the year in order to confirm these findings.     

Cilia-associated respiratory (CAR) bacillus infection in adult red deer, chamois, and roe deer

Cilia-associated respiratory (CAR) bacillus is an unclassified bacterium that colonizes the ciliated epithelium of airways in laboratory rats, laboratory mice, and laboratory and conventionally reared rabbits, cattle, goats, and pigs. Data on the prevalence of CAR bacillus infection in wild animals are lacking. The present study demonstrated the occurrence of the organism in wild red deer (Cervus elaphus hippelaphus), chamois (Rupicapra rupicapra), and roe deer (Capreolus capreolus) from the Val Fontana in northern Italy. Prevalence ranged from 26% for red deer to 56% for chamois, with a statistically significant negative correlation between CAR bacilli infection and the presence of lymphoid follicles.     

Urinary cortisol and urea nitrogen responses in irreversibly undernourished mule deer fawns

We examined the concentration of urinary cortisol and urea nitrogen of five hand-reared mule deer (Odocoileus hemionus) fawns that failed to recover from winter starvation, and compared them to levels found in fawns that recovered. The fawns wintered in fenced pastures stocked with wild deer, and were put back on supplemental feed after losing 15% of their body mass. The five fawns that died began receiving supplemental feed up to 3 wk before death. All continued to lose weight, and were consequently removed from the pasture and fed ad libitum 4 to 10 days before death. In the animals that died, cortisol levels continued to increase regardless of food availability, and were correlated with those of urea nitrogen. Postmortem cortisol and urea nitrogen measurements were significantly greater than concentrations found in the weeks preceding death. We hypothesize that uncontrolled protein catabolism is promoted by high levels of cortisol. These cortisol levels may reach a point at which irreversible multiple-system organ failure occurs, leading to the animal's death.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

Numerical responses of an avian predator to prey fluctuations in a temperate latitude: breeders vs. entire population

Distinct numerical responses of predators to fluctuations in the abundance of their prey are often observed in northern regions but occur more rarely in temperate latitudes. This statement is, however, mostly based on observations of breeding populations, while in some predators, for example in raptors, numerous non-breeding floaters can occur. I estimated the breeding density and reproductive performance (nest survey) as well as the density of entire population (transects with distance sampling) of the common buzzard Buteo buteo in western Poland (52°N) in the years 2005–2014 to test the hypotheses that in temperate latitudes: (1) the breeding population of these birds does not show any numerical response, understood as annual changes in their abundance; (2) its reproductive success, however, changes with the abundance of main prey, the common vole Microtus arvalis; and (3) hence the entire buzzard population (including potential immature floaters) present in a given area during the nesting period responds numerically with some time delay. The reproductive success of buzzards was positively correlated with their prey abundance. Contrary to my predictions, however, the breeding population of buzzards showed a slight numerical response with a 3-year lag and the entire population tracked vole fluctuations without any time delay. The immediate numerical response of the entire buzzard population was probably caused by large-scale movements of floaters. Such rapid numerical responses of some predators may contribute to the relative stability of prey populations in temperate latitudes compared to northern regions.     

Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland

A pattern of wolfCanis lupus Linnaeus, 1758 predation on red deerCervus elaphus Linnaeus, 1758 was studied in Bieszczady Mountains in 1991–2002. In total 324 remains of red deer > 4 months old, killed by wolves throughout the year, were found. The sex, age and bone marrow fat content of wolf kills were compared with the same characteristics within the free living red deer population. The overall contribution of calves killed by wolves (24%) in October-May was higher than in the population (17%), and decreased from autumn to spring. Adult males were more vulnerable to wolf predation than adult females: stags constituted 62% and hinds 38% of adult red deer killed by wolves, whereas in the population, the percentages were 37 and 63%, respectively. Stags killed by wolves were younger ([`(x)] = 4.1\bar x = 4.1 years old) than hinds ([`(x)] = 8.9\bar x = 8.9 years old). Wolves killed more > 8 years old hinds and < 5 years old stags than available in the population. In wolf kills, the average fat content in femur marrow was higher among hinds (84.9%) than stags (69.3%) and calves (66.1%). Only 8% of hinds had < 70% femur marrow fat content, whereas 40% of calves and 38% of stags had marrow fat values below that level. Marrow fat content showed seasonal variation and was the lowest in March among all sex-age classes. The monthly share of stags in all kills, and hinds in hind-calf part of the sample was negatively correlated with their monthly average bone marrow fat content, and monthly share of calves was positively correlated with monthly average bone marrow fat content of adults. The segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter until spring contribute to the high overall incidence of calves and adult males and the relatively low incidence of adult females among wolf kills.     

Invading parasites cause a structural shift in red fox dynamics

The influence of parasites on host life histories and populations is pronounced. Among several diseases affecting animal populations throughout the world, sarcoptic mange has influenced many carnivore populations dramatically and during the latest epizootic in Fennoscandia reduced the abundance of red fox by over 70%. While the numerical responses of red fox populations, their prey and their competitors as well as clinical implications are well known, knowledge of how sarcoptic mange affects the structure of the dynamics of red fox populations is lacking. Integrating ecological theory and statistical modelling, we analysed the long-term dynamics (1955-1996) of 14 Danish red fox populations. As suggested by the model, invading sarcoptic mange significantly affected direct and delayed density dependence in red fox dynamics and concomitant shifts in fluctuation patterns were observed. Our statistical analyses also revealed that the spatial progressive spread of mange mites was mirrored in the autocovariate structures of red fox populations progressively exposed to sarcoptic mange.     

Autumn-winter diet overlap of fallow, red, and roe deer in forest ecosystems, Southern Poland

The wild population of fallow deer in Central Europe has grown considerably over the last decade. However, information on feeding habits of this alien species in relation to the indigenous red deer or roe deer, in areas of their co-occurrence, is scarce. A prevailing view maintains that their food-niches are distinct, although direct comparative studies have not been carried out. Therefore, the aim of the research was to compare the diets of fallow, red, and roe deer feeding in the same habitat. Research was based on the rumen contents of 242 animals hunted in the autumn-winter season in the forests of Southern Poland. The analyses demonstrated that fallow deer are moderate grazers in such conditions and eat more graminoids in comparison to red or roe deer (36.4% vs. 16.1% or 5.5%, respectively). On the other hand, it feeds on less browse (17.2% vs. 41.4%) or dwarf shrubs (8.4% vs. 19.0%) than red deer, and on less bramble (10.9% vs. 34.6%) or forbs (4.0% vs. 7.6%) in comparison to roe deer (P=0.05). Although the diets of the three deer species differ in terms of the proportion of each food type in their diet, overlapping of their food-niches is high (52.6%).     

Variation in maternal investment in a small cervid; the effects of cohort,sex, litter size and time of birth in roe deer (Capreolus capreolus) fawns

 We investigated the effects of cohort, sex, litter size and time of birth on birth weights and postnatal growth rates of roe deer fawns in a highly reproductive Norwegian population. By repeatedly recapturing radio-collared individuals, a total of 950 weights were obtained from 231 fawns of known age. In accordance with earlier studies, there was a period of linear growth during the first month following birth. Mean postnatal growth rates of 155 g/day are the highest yet recorded for roe deer; however, the mean birth weights of fawns were lower than those reported from populations in continental Europe. During the period of linear growth, we found no sex differences. However, growth rates were affected both by time of birth and litter size; fawns born early had lower growth rates than fawns born during or after the peak calving period, and fawns in triplet – groups had lower growth rates than either fawns in twin – groups or single fawns. Despite a fourfold increase in population density during the study, this factor was not able to explain variation in postnatal growth rates, although cohort effects on birth weight were evident. Received: 13 May 1996 / Accepted: 26 June 1996     

Fast and efficient DNA-based method for winter diet analysis from stools of three cervids: moose, red deer, and roe deer

Effects of cervid browsing on timber production, especially during winter, lead to economic losses in forest management. The aim of this study was to present an efficient DNA-based method which allows qualitative assessment of the winter diet from stools of moose (Alces alces), roe deer (Capreolus capreolus), and red deer (Cervus elaphus). The preliminary results of the diet composition of the three cervids from Poland were also presented with a special emphasis on moose. The electropherograms of the chloroplast intron trnL (UAA) P6 loop amplification products using g (fluorescence-labeled) and h primers revealed differences in the length of PCR products among various plant species eaten by these herbivores. In addition, the usage of species-specific primers allowed unambiguous identification of different gymnosperms and angiosperms. The preliminary moose diet analysis, based on winter fecal samples from the entire range of moose occurrence in Poland, revealed the presence of 15 to 24 tree, shrub, and herbaceous species. This fast, cost-efficient, and simple method proved also to be reliable for the diet analysis of red deer and roe deer. It may be a valuable tool in forest and conservation management, as well as a way of enhancing ecological studies focusing on the impact of herbivores on the ecosystems and their possible food niche overlap.