The occurrence of hepatic trematodes in sheep from Emilia-Romagna

In the present investigations, 563 samples of ovine faeces from 32 flocks of Emilia-Romagna have been examined for the presence of hepatic trematodes eggs. All flocks proved positive for Dicrocoelium dendriticum and 16 for Fasciola hepatica; 511 (90,76%) out of 563 animals proved positive, with 507 (90,05%) for D. dendriticum and 50 (8,88%) for F. hepatica, 46 of these last ones were positive for both parasites.     

The use of MM3 monoclonal antibodies for the early immunodiagnosis of ovine fascioliasis

This study reports a new capture ELISA (MM3-SERO) for the serodiagnosis of sheep fascioliasis, based on the use of the monoclonal antibody (mAb) MM3. Like our previously reported indirect ELISA method, based on the use of a FPLC-purified fraction (fraction IV) of the Fasciola hepatica excretion/secretion antigens (ESAs), this new test was able to detect animals infected with very small numbers of metacercariae (5-40) and showed no cross-reaction with sera from sheep infected with other parasites, i.e., Moniezia spp., Cysticercus tenuicollis, and Dicrocoelium dendriticum. In contrast with these 2 methods, some sera (mainly those obtained from animals infected with D. dendriticum) showed high reactivities in indirect ELISA with whole F. hepatica ESAs used as control. Interestingly, the MM3-SERO ELISA has a better signal-to-noise ratio than the fraction-IV ELISA, thus allowing detection of seroconversion in infected sheep on average 1 wk earlier (3.2 +/- 0.4 wk postinfection [PI] for MM3-SERO ELISA vs. 4.2 +/- 0.9 wk PI for fraction IV ELISA). Moreover, the antibody response detected with MM3-SERO ELISA was more uniform, with seroconversion always occurring at 4 wk PI in sheep with 1-2 flukes and at 3 wk PI in sheep with more than 2 flukes. The MM3-SERO ELISA was also used to evaluate the kinetics of antibody response against MM3-recognized antigens in sera from sheep experimentally infected with F. hepatica and then treated with triclabendazole. Our results showed that antibody levels dropped by about 25% during the 4-wk observation period following the flukicide treatment, whereas they remained invariably high in all sheep left untreated. We conclude that the MM3-SERO ELISA is a 100% sensitive and 100% specific test for the early serodiagnosis of sheep fascioliasis. Preliminary studies in our laboratory seem to indicate that this method may also be useful for the determination of anti-F. hepatica antibodies in serum and milk of other ruminants. A commercial version of MM3-SERO is currently available from BIO X Diagnostics (La Jemelle, Belgium).     

Isoenzymes of lactate dehydrogenase (EC 1.1.1.27) in Dicrocoelium dendriticum and Fasciola hepatica (Trematoda)

The isoenzymes of LDH (EC 1.1.1.27) were studied in Dicrocoelium dendriticum and Fasciola hepatica by horizontal electrophoresis on polyacrylamide gel. Six isoenzymes in D. dendriticum and four in F. hepatica were detected. Densitometric scans demonstrated that the enzyme pattern of LDH in D. dendriticum parasitizing hepatic tissue of Capra hircus was clearly different from the one obtained when the trematode parasitized other hosts such as Bos taurus or Ovis aries.     

Parasiticidal effects of peroxynitrite on ovine liver flukes in vitro

Peroxynitrite (ONOO-) is a cytotoxic anion, produced by interaction between nitric oxide and superoxide in vivo in some inflammatory cells. This study investigated its effects on Fasciola hepatica and Dicrocoelium dendriticum isolated from ovine livers and kept in bile at room temperature. Peroxynitrite was synthesized using a quenched flow reactor and assayed spectrophotometrically. It was applied at different concentrations (10(-3.5) to 10(-2.3 M)) to the flukes kept in bile. The viability of the peroxynitrite-treated flukes was compared with a control group (n=5-7 per group). Control F. hepatica and D. dendriticum lived for 226+/-11and 208+/-14min, respectively. Life times were decreased by peroxynitrite at all concentrations used (P<0.001). At the highest concentration of peroxynitrite, F. hepatica and D. dendriticum lived only for 6.1+/-0.4 and 4.1+/-4.1+/-0.2min, respectively. Correlation between peroxynitrite concentration and parasite viability was significant in the case of F. hepatica (r= -0.842; P= 0.0035). A single application of peroxynitrite can decrease the life span of ovine liver flukes. A failure in the activation of hepatic macrophages in infected animals may lead to a decreased production of free radicals and, thus, peroxynitrite. Such a failure is likely to deprive the body of a defence tool against multicellular parasites.     

Arthropod and helminth parasites of the wild guinea pig,Cavia aperea,from the Andes and the cordillera in Peru,South America

As part of an ongoing research project concerning the diversity and distribution of parasites of Caviidae in South America, 143 wild guinea pigs (Cavia aperea) were collected from 3 localities in the Andean Highlands of Peru. Samples were collected between November 1996 and May 1999 and included representatives of arthropods, nematodes, and trematodes. Seven species of arthropods: Leptopsylla segnis (27.3%), Tiamastus cavicola (6.9%), Gliricola porcelli (55.2%), Hoplopleura alata (6.9%), Polyplax spinulosa (12.6%), Myobia musculi (1.4%), and Eutrombicula bryanti (49.6%); 4 species of nematodes: Capillaria hepatica (6.9%), Graphidioides mazzai (18.8%), Trichuris gracilis (3.5%), and Paraspirudera uncinata (37%); and a single trematode, Fasciola hepatica (4.2%), were identified.     

Bovine viral diarrhea virus multiorgan infection in two white-tailed deer in southeastern South Dakota

The susceptibility of wild ruminants, especially cervids, to bovine viral diarrhea virus (BVDV) has remained an enigma. Two white-tailed deer (Odocoileus virginianus) were submitted to the Animal Disease Research and Diagnostic Laboratory (ADRDL) in the fall of 2003 by the South Dakota Game Fish and Parks for chronic wasting disease (CWD) testing. Both animals were CWD negative. The animals were necropsied and histopathology, viral antigen detection, and virus isolation were performed. A noncytopathic (NCP) BVDV was isolated from the lungs and several other tissues of both animals. Formalin-fixed ear notches from both animals were positive for BVDV antigen by immunohistochemistry. The BVDV isolates were typed with the use of polymerase chain reaction in 5' untranslated region (UTR) and one isolate was typed a Type 2a and the other a Type 1b. Future field surveys to determine the incidence of BVDV along with experimental studies to determine if white-tailed deer fawns can be persistently infected with BVDV are needed.     

An ultrasensitive capture ELISA for detection of Fasciola hepatica coproantigens in sheep and cattle using a new monoclonal antibody (MM3)

A capture enzyme-linked immunosorbent assay (ELISA) using a new monoclonal antibody (mAb MM3) is reported for the detection of Fasciola hepatica excretory-secretory antigens (ESAs) in feces of infected hosts. The mAb MM3 was produced by immunization of mice with a 7- to 40-kDa purified and O-deglycosylated fraction of F. hepatica ESAs, which has previously been shown to be specific for the parasite. The specificity and sensitivity of the MM3 capture ELISA were assessed using feces from sheep and cattle. Sheep feces were obtained from a fluke-free herd (with most animals harboring other nematodes and cestodes), from lambs experimentally infected with 5-40 F. hepatica metacercariae and in some cases treated with triclabendazole at 14 wk postinfection (PI), and from uninfected control lambs. Cattle feces were collected at the slaughterhouse from adult cows naturally infected with known numbers of flukes (from 1 to 154) or free of F. hepatica infection (though in most cases harboring other helminths). The MM3 capture ELISA assay had detection limits of 0.3 (sheep) and 0.6 (cattle) ng of F. hepatica ESA per milliliter of fecal supernatant. The assay detected 100% of sheep with 1 fluke, 100% of cattle with 2 flukes, and 2 of 7 cattle with 1 fluke. The false-negative animals (5/7) were probably not detected because the F. hepatica individuals in these animals were immature (5-11 mm in length). As expected, coproantigen concentration correlated positively (r = 0.889; P < 0.001) with parasite burden and negatively (r = 0.712; P < 0.01) with the time after infection at which coproantigen was first detected. Nevertheless, even in animals with low fluke burdens (1-36 parasites), the first detection of F. hepatica-specific coproantigens by the MM3 capture ELISA preceded the first detection in egg count by 1-5 wk. In all sheep that were experimentally infected and then untreated, coproantigen remained detectable until at least 18 wk PI, whereas in sheep that were experimentally infected and then flukicide treated, coproantigen became undetectable from 1 to 3 wk after treatment. None of the fecal samples from sheep or cattle negative for fascioliasis but naturally infected with other parasites including Dicroelium dendriticum showed reactivity in the MM3 capture ELISA. These results indicate that this assay is a reliable and ultrasensitive method for detecting subnanogram amounts of F. hepatica antigens in feces from sheep and cattle, facilitating early diagnosis.     

Forestomach pH in hunted roe deer (Capreolus capreolus) in relation to forestomach region, time of measurement and supplemental feeding and comparison among wild ruminant species

There is a debate whether supplemental feeding of deer bears the risk of inducing health problems, in particular acidosis. Here, the pH values of forestomach contents of free-ranging roe deer (Capreolus capreolus) shot in areas with and without supplemental winter feeding were compared. pH was similar in the dorsal and ventral rumen, but lower at these sites than in the Atrium ruminis, where it was again lower than in the reticulum; this pattern corresponds to expectations based on differences in the presence of saliva at the different sites of the forestomach. pH was lower with increasing time that elapsed between death of the animal and measuring pH in unsupplemented animals and was lower in unsupplemented animals in May/June than later in the year. Animals with supplemental winter feeding had significantly lower rumen pH (5.5) than animals without food supplementation (5.7). These data suggest that supplemental feeding of roe deer has the potential to lower forestomach pH. Although pH values measured in supplemented animals in this study would be considered indicative of rumen acidosis in domestic cattle, they are within the range previously measured in various free-ranging Odocoilid species, including roe deer; were of a similar magnitude as the May/June values of unsupplemented roe deer in this study; and must be considered with respect to potentially rapid declines in pH between death of the animal and pH measurement. Given methodological problems, analyses of literature data from free-ranging wild ruminants provide little evidence for a systematic variation of rumen pH with feeding type and body mass, but lead to the hypothesis that some New World cervids, including the roe deer, might either naturally have lower pH values than other ruminants or rumen contents whose pH drops rapidly after death.     

Cortisol response of wild ungulates to trauma situations: hunting is not necessarily the worst stressor

Animal welfare concerns are becoming a central issue in wildlife management and conservation. Thus, we investigated stress response of wild ungulates to potentially traumatic situations (shooting injuries, vehicle collisions, entanglement, injuries or diseases) and hunting methods (stalking, battues and hunts with dogs) by means of serum cortisol concentrations from blood collected from killed animals. Cortisol levels in roe deer ranged below and in wild boar above levels for moose, red deer and fallow deer (hence, pooled as a group “deer”). Apart from species, cortisol concentration in trauma situations was mainly explained by trauma type and presence of disturbance after the trauma event. Effect of trauma type differed significantly for “deer”, with animals caught in fences and suffering vehicle collisions experiencing higher cortisol levels than animals injured by shooting. Differences between hunting methods were observed in the cervids (“deer” and roe deer), with stalking leading to lower cortisol levels than hunts with dogs (both groups) and battues (roe deer). Events both before and after the shot, such as duration of pursuit prior to shooting, location of injury, trauma length and presence of disturbance after the shot were relevant for cortisol levels in hunted cervids. Our results indicate that search teams tracking and euthanising wounded animals should behave in a calm way to minimise disturbance. Still, it is important to acknowledge that many situations described in the literature, i.e. reindeer handling, roe deer captures and red deer yarding, seem even more stressful, beside vehicle collisions, than most hunting methods.     

Bluetongue epidemiology in wild ruminants from Southern Spain

Serum samples from 210 wild ruminants collected between 2006 and 2007 in southern Spain were tested for antibodies against bluetongue virus (BTV) by means of a competitive ELISA assay. Eighty-seven of the 210 wild ruminants analysed (41%) showed antibodies against BTV. Statistically significant differences were found in the seroprevalence among species: 66% (65 of 98) for red deer (Cervus elaphus), 50% (ten of 20) for fallow deer (Dama dama), 33% (three of nine) for mouflon (Ovis aries musimon) and 11% (nine of 83) for Spanish ibex (Capra pyrenaica). Overall, the sites where seropositive wild ruminants were found coincide with the areas where BTV had been detected in livestock, but in eastern Sierra Morena, the virus circulated in wild ruminants, although it had not been detected in domestic ruminants in the same areas. Wild ruminants over 1-year of age (sub-adults and adults) had significantly higher seroprevalences than juvenile animals. Statistically significant differences were also observed between BTV seroprevalence and management (free-ranging vs. captivity) with higher prevalence in free-ranging animals. The high seroprevalences obtained suggest that BTV is widespread in wild ruminants in southern Spain. This factor could have an important influence on the evolution of the infection in domestic livestock and indicates the need to include wild ruminant species in BTV surveillance or control programs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fence-Line Contact Between Wild and Farmed Cervids in Colorado: Potential for Disease Transmission

Abstract: Direct and indirect contact between wild and farmed cervids along perimeter fences may play a role in transmission of diseases like chronic wasting disease (CWD), but no studies have quantified such interactions. At 9 high-fenced commercial elk (Cervus elaphus) farms in Colorado, USA, during October 2003 to January 2005, we used animal-activated video to estimate rates of fence-line use by wild cervids, rates of direct contact between farmed and wild cervids, and probability of direct contact when wild cervids were present. We recorded 8-fold-more wild elk per unit time than mule deer (Odocoileus hemionus) at fence lines. Depending on site, we recorded 0.66 to 46.90 wild elk per 1,000 hours of camera monitoring. We documented 77 interactions between wild and farmed elk involving naso-oral contact and no contact between wild mule deer and farmed elk. Rate of direct contact ranged from 0.00 to 1.92 direct contacts per 1,000 hours of camera monitoring among sites. Given recorded presence of wild elk, estimated probability of observing direct contact during a 2-minute video recording ranged from 0.00 to 0.11 among sites. Risk of direct contact was about 3.5 times greater for single woven-wire fence compared with offset electric fence attached to a single woven-wire fence. We observed no direct contact through double woven-wire fences. Because little is currently known about infection rates associated with infection mechanisms, we cannot infer a level of CWD infection risk from our results, but some form of double fencing should reduce potential for direct and indirect transmission of disease into or out of elk farms.     

The helminth fauna of wild and domestic ruminants in Mongolia—a review

The helminth fauna of eight species of wild ruminants and five species of domestic ruminants sharing common grassland in Mongolia was reviewed. A total of 108 species of helminths, belonging to 40 genera, 23 families, 11 superfamilies, 6 orders and 3 classes were reported in wild and domestic ruminants in Mongolia. Several new host–helminth associations have been recorded. Twenty seven helminth species have been found in certain host species for the first time in Mongolia. These included 2 species of trematodes, 2 species of cestodes and 23 species of nematodes. Helminths that are specific for some of the examined ruminant species were determined. However, wild animals may serve as a reservoir of helminth infections for domestic animals when sharing grassland in Mongolia. The helminth fauna of the ruminants with respect to occurrence, distribution, prevalence and intensity of infection was considered for six geographic regions of Mongolia.     

Evidence of <Emphasis Type="Italic">Anaplasma phagocytophilum</Emphasis> and <Emphasis Type="Italic">Rickettsia helvetica</Emphasis> infection in free-ranging ungulates in central Slovakia

The purpose of this study was to investigate the role of wild animals for Anaplasma phagocytophilum, other ehrlichiae/anaplasmae, Rickettsia helvetica and other rickettsiae and whether different genetic variants of A. phagocytophilum in central Slovakia exist. A total of 109 spleen samples from 49 red deer (Cervus elaphus), 30 roe deer (Capreolus capreolus), 28 wild boar (Sus scrofa) and two mouflon (Ovis musimon) were collected from June 2005 to December 2006. Polymerase chain reaction (PCR) amplification of the16S rRNA gene was used for detection of ehrlichiae/anaplasmae. A nested PCR targeting part (392 bp) of groESL gene was applied for the specific detection of A. phagocytophilum. Fragments of the gltA and ompA genes (381 bp and 632 bp, respectively) were amplified to detect rickettsiae, followed by sequencing. A. phagocytophilum and R. helvetica were detected in wild animals. The prevalence of A. phagocytophilum was 50.0 ± 18.2% in roe deer and 53.1 ± 14.1% in red deer. None of the 28 wild boar was PCR positive for ehrlichiae/anaplasmae. A. phagocytophilum was detected in one mouflon. R. helvetica was found in one roe deer. Our study suggests a role of cervids as a natural reservoir of A. phagocytophilum in Slovakia. However, the role of cervids and wild boars in the circulation of R. helvetica remains unknown. The analysis of sequence variation in the msp4 coding region of A. phagocytophilum showed the presence of different variants previously described in ruminants.     

Malignant catarrhal fever in free-ranging cervids associated with OvHV-2 and CpHV-2 DNA

Pathologic lesions were summarized in 18 free-ranging cervids (15 moose [Alces alces], two roe deer [Capreolus capreolus], and one red deer [Cervus elaphus]) diagnosed with malignant catarrhal fever (MCF) after examination at the National Veterinary Institute, Oslo 1982-2005. Eye lesions (conjunctivitis, corneal opacity, fibrin clots in the anterior eye chamber) were the most frequent gross finding. Erosive-ulcerative mucosal lesions in the nose and mouth were also commonly found. Histopathology revealed a nonpurulent vasculitis and perivasculitis in the central nervous system (CNS) typical of MCF in 16 of the cases. The diagnosis in the remaining two animals was based upon histologic eye lesions consistent with MCF (CNS not available for examination). Polymerase chain reaction was run on samples from 15 individuals for evidence of MCF-virus DNA, and ovine herpesvirus-2 (OvHV-2) DNA was detected in five moose, one roe deer, and one red deer, and caprine herpesvirus-2 (CpHV-2) DNA was detected in two moose and one roe deer. Sera from 1,000 free-ranging cervids were tested for specific antibodies to MCF-associated viruses (MCFV) by competitive inhibition enzyme-linked immunosorbent assay. The seroprevalences were: red deer 5%, reindeer (Rangifer tarandus) 4%, roe deer 2%, and moose 0.4% (n = 250 for all four species). The results indicate that sheep and goat MCFV may cause serious disease in wild moose, roe deer, and red deer. The seropositive cervids most likely represent individuals infected with either OvHV-2 or CpHV-2, but may also reflect infections with other related MCFV.     

Seroepidemiology of chlamydial infections of wild ruminants in Spain

Chlamydial infections were determined serologically among wild ruminants in the Nature Park of the Sierras de Cazorla, Segura y Las Villas (CNP; Spain). Sampling was done during the period from 1990-95. There were 1,244 blood samples collected, consisting of 490 from fallow deer (Dama dama), 343 from mouflon (Ovis mussimon), 283 from red deer (Cervus elaphus) and 128 from Spanish ibex (Capra pyrenaica). Specific complement-fixing antibodies of Chlamydia spp. were detected by means of microtechnique, using lipopolysaccharide antigen. The relationship of biological (species, sex, age), temporal (year) and territorial (central and peripheral areas) factors to seropositive prevalence was examined, and preliminary data were collected on whether or not sheep and goat herds grazing in the peripheral areas of the park also were infected with Chlamydia spp. Chlamydiosis was common in the four species of wild ruminants in the CNP in all the years studied. The prevalence of Chlamydia sp. in mouflon (37%) was significantly greater than in fallow deer (30%), and both had a significantly higher prevalence rate than Spanish ibex and red deer (both 24%). The four species of wild ruminants were similar in that they act as reservoirs of Chlamydia spp., although their receptivity may be different, and the infection can certainly be maintained among these animals by intra-group transmission. The differences in prevalences and geometric mean titers (GMT), both between the sexes (male versus female) and between different ages (adult versus juvenile), were insignificant in all four species. For all species of wild ruminants both prevalence rates and GMTs were greater in populations occupying the peripheral areas of the park than in those inhabiting the central area. Herds of sheep and goats had a high prevalence of chlamydiosis. Intertransmission of Chlamydia sp. between wild and domestic ruminants occurred through grazing on the same pastures. The highest mean prevalence (44%) of patent infections (CFT titers of > or =1:80) was detected in red deer, although this frequency was not significantly different from those observed in mouflon (39%), Spanish ibex (38%), and fallow deer (37%). The proportion of patent infection was higher in females than in males, and none of the juveniles (<2-yr-old) showed patent infections. The prevalence of predicted patent chlamydial infections was always higher in the peripheral areas of the park, although only among mouflon and fallow deer were the differences statistically significant.     

Patterns of antler consumption reveal osteophagia as a natural mineral resource in key periods for red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)

Osteophagia is a common behaviour among ungulates. It has been considered as a mineral source for cervids, mainly related to phosphorus deficiency. In this study, we aimed to study the seasonality in antler consumption of Iberian red deer (Cervus elaphus) and to assess the differences among sex/age classes. In a game reserve located in South-Eastern Spain, antlers were offered to free-ranging animals, and their chewing behaviour was recorded with camera traps. Red deer was the species observed with greatest frequency (90.5%), and only red deer was observed chewing the offered antlers. Males showed greatest peak of consumption at the end of antler growth; females showed the greatest peak at the beginning of the lactation and calves after delivery of the newborn. All groups showed an important second peak of consumption in September. We conclude that osteophagia might have a role as a natural mineral supplement for red deer, but not for wild boar or foxes; and that each sex/age class uses this resource at different times according to differential needs in calcium and phosphorus, supporting the “seasonally increased requirements hypothesis”.     

Economical strategies in the Upper Palaeolithic in the Cantabrian region

The aim of this paper is to survey the economic approach in relation to animal use during Upper Palaeolithic times in Northern Spain. The animals present in the archaeological record can be separated into main groups, like those present by natural reasons and those introduced by human groups. Also in the anthropic aported animals, some of them are related to food procurements and others to economic reasons. Animals like deer, bovids, equids and so on, were mainly related to nutrition, while some others like mustelids, canids or felines were more related to the obtention of furs or other non-alimentary uses. The analysis of the “main-six” (red deer, roe deer, big bovids, horse, ibex and wild boar) showed that all of them were used throughout the Upper Palaeolithic without many changes in time.     

Deaths from exertional myopathy at the National Zoological Park from 1975 to 1985

A retrospective study was conducted to determine the incidence of deaths from exertional myopathy (EM) in the hoofstock collection at the National Zoological Park (NZP) from 1975 to 1985. The diagnosis of EM was based on history, clinical signs, clinicopathological findings, gross and microscopic lesions. Only bovids, cervids, and equids were included in the study. Data were derived from the medical and pathological records of the NZP. There were 10 cases of EM during this period, but only seven deaths were directly attributable to EM. Five deaths occurred after immobilization. Two deaths occurred after improper transport, but the anesthetic history was not known in these cases. Pathologic changes consistent with EM were incidental findings in an animal killed by dogs, and in two animals euthanized because of suspected infection with Parelaphostrongylus tenuis. Only cervids and bovids were affected. There was no age or sex predilection for developing EM and deaths occurred throughout the year. Deaths associated with immobilization occurred most often after anesthesia with xylazine and etorphine. These are the drugs used most frequently in bovid and cervid anesthesia at the NZP. The overall incidence of deaths from EM after immobilization was 0.25% for the 10-yr period.     

Abomasal parasites in wild sympatric cervids, red deer, Cervus elaphus and fallow deer, Dama dama, from three localities across central and western Spain: relationship to host density and park management

A survey of abomasal parasites in cervids from Central Spain was conducted at 3 sites, Quintos de Mora (Toledo), Malué?ez de Arriba (Cáceres), and La Herguijuela (Cáceres). Commonly occurring helminths belonged to 3 polymorphic species of the Ostertagiinae: Spiculopteragia asymmetricalS. quadrispiculata, Ostertagia leptospicularis/O. kolchida, and O. drozdzi/O. ryjikovi. Trichostrongylus axei was found in very few cases. Ostertagia drozdzi/O. ryjikovi and the minor male morphotype, S. quadrispiculata, are reported for the first time in red deer from Spain. The 3 ostertagiine species are also reported for the first time in fallow deer from Spain. These 3 species of Ostertagiinae are primarily parasites of cervids, and nematode species characteristic of domestic ruminants were not present. Prevalence of infection by gastrointestinal parasites in cervids was high, ranging from 97.5 to 100%, across the 3 areas sampled. Mean intensity of infection and abundance showed a positive relationship to the population density of red deer. Helminth burdens were higher in fallow deer than in the sympatric red deer and may reflect the gregarious social structure and different foraging patterns of fallow deer.