Tick-borne parasitic diseases in cattle: current knowledge and prospective risk analysis related to the ongoing evolution in French cattle farming systems

Parasitic diseases, like babesiosis and theileriosis are transmitted by ticks: their occurrence is therefore linked to the size of the tick stock and the seasonality of the vectors. Babesia divergens bovine babesiosis transmitted by Ixodes ricinus is widespread and often reported in France. Serological prevalence is high, ranging from 20 to 80% according to the farms. Clinical incidence is low: around 0.4% for the whole cattle population. The endemic situation is unstable and clinical cases occur more frequently with farming system modifications. L. ricinus is a tick essentially found in woodlands and so, for the most part, is found in closed rural areas. The situation of the other bovine babesiosis (Babesia major) and theileriosis (Theileria orientalis) is not well documented in France. However, the epidemiology of parasitic diseases is changing, especially because of changes in the environmental characteristics, i.e. both farm and herd management conditions and also climatic conditions. These modifications can provoke an increase in the tick stocks, an increase in the contact rate between cattle and ticks, and an increase in the contact rate between cattle and the wild fauna, especially deer. This results in likely modifications of the endemic situation, with a higher risk of clinical babesiosis in the medium term.     

Ticks on livestock in Barbados

Cattle, sheep, goats, and horses in most areas of Barbados were examined for ticks. About 68% of the cattle, mostly Holstein dairy cows, and 23% of the sheep, mostly purebred or crossbred black belly sheep, were infested with the southern cattle tick, Boophilus microplus (Canestrini), a vector of bovine babesiosis and anaplasmosis. About 13% of the horses were infested with the tropical horse tick, Anocentor nitens (Neumann), a vector of equine babesiosis. Cattle owners used a variety of acaricides as dusts, sprays, or washes on their cattle for the control of ticks.     

Tick control methods used by resource-limited farmers and the effect of ticks on cattle in rural areas of the Eastern Cape Province,South Africa

A survey to document tick control methods used by resource-limited farmers in the control of cattle ticks in the Eastern Cape Province of South Africa was conducted by interviewing 59 cattle farmers using structured questionnaires and general conversation. Information collected was on external parasites of cattle, their effects and their control methods. Ticks were reported to be a major problem causing diseases like anaplasmosis (89.8%), babesiosis (55.9%) and ehrlichiosis (16.9%), as well as wounds that predispose to screwworm infestation, tick worry and teat damage to cows troubling farmers in their farming enterprises. The main tick control methods were; acaricides provided by government, however 94.9% of the farmers interviewed were of the opinion that the dip wash is not effective in killing the ticks. As a result, farmers complement the government dipping service with their own initiatives like spraying with conventional acaricides (22%), household disinfectants such as Jeyes fluid (18.6%), used engine oil (10.2%), chickens (5.1%), manual removal (5.1%), and pouricides (1.7%). In addition, some farmers also use plants (6.8%), mainly the leaf of Aloe ferox and the bark of Ptaeroxylon obliquum. The study revealed ticks to be a major problem in the study area.     

Disease at the wildlife-livestock interface: Acaricide use on domestic cattle does not prevent transmission of a tick-borne pathogen with multiple hosts

Several prominent and economically important diseases of livestock in East Africa are caused by multi-host pathogens that also infect wildlife species, but management strategies are generally livestock focused and models of these diseases tend to ignore the role of wildlife. We investigate the dynamics of a multi-host tick-borne disease in order to assess the efficacy of tick control from an ecological perspective. We examined the efficacy of a widespread measure of tick control and developed a model to explore how changes in the population of ticks due to control measures on cattle impact dynamics of Theileria parva infection in a system with two primary host species, cattle and Cape buffalo (Syncerus caffer). We show that the frequency of acaricide application has a significant impact on the tick population both on the host and in the environment, which can greatly reduce the pathogen load in cattle. We also demonstrate that reducing the tick population through cattle-related control measures is not sufficient to diminish disease transmission in buffalo. Our results suggest that under current control strategies, which target ticks on cattle only, T. parva is likely to remain a significant problem in East Africa, and require the continued use of acaricides, which has significant economic and ecological consequences.     

Babesias of red deer (Cervus elaphus) in Ireland

ABSTRACT: Blood samples were obtained from 38 wild red deer (Cervus elaphus) at two sites in Ireland and subjected to PCR analysis of the 18S rRNA gene followed by sequencing. Two fragments of the 18S rRNA gene were generated by two different PCR protocols and subsequent sequencing suggested that at least six of the deer were infected by a babesia that, in those loci, is indistinguishable from Babesia divergens, an important tick-borne pathogen of cattle and of zoonotic significance. Additionally, a B. odocoilei-like parasite was detected in three samples and a babesia that did not match any sequences in the GenBank database was found in five samples. Neither B. capreoli nor B. venatorum (EU1) were found. There have been several reports of B. divergens occurring in deer species, including red deer, roe deer (Capreolus capreolus) and reindeer (Rangifer tarandus). However, in view of recent re-sequencing of bovine-origin samples deposited previously in GenBank, it is unlikely that any of these sequences from deer are B. divergens. The present study describes the only deer piroplasm detected so far that shows complete identity with B. divergens, in just over half of the 18S rRNA gene. The entire gene of this deer parasite should be analysed and transmission experiments undertaken before the infectivity of B. divergens for red deer can be confirmed.     

Molecular identification of tick-borne pathogens in Nigerian ticks

A molecular epidemiology investigation was undertaken in two Nigerian states (Plateau and Nassarawa) to determine the prevalence of pathogens of veterinary and public health importance associated with ticks collected from cattle and dogs using PCR, cloning and sequencing or reverse line blot techniques. A total of 218 tick samples, Amblyomma variegatum (N=153), Rhipicephalus (Boophilus) decoloratus (N=45), and Rhipicephalus sanguineus (N=20) were sampled. Pathogens identified in ticks included piroplasmids (Babesia spp., Babesia bigemina and Babesia divergens), Anaplasma marginale and Rickettsia africae. Piroplasmids were identified in A. variegatum, A. marginale was found in R. decoloratus, while R. africae was detected in all tick species examined. Ehrlichia spp. and Theileria spp. were not identified in any of the ticks examined. Of the 218 ticks examined, 33 (15.1%) contained pathogen DNA, with the presence of B. divergens and R. africae that are zoonotic pathogens of public health and veterinary importance. The variety of tick-borne pathogens identified in this study suggests a risk for the emergence of tick-borne diseases in domestic animals and humans, especially amongst the Fulani pastoralists in Plateau and Nassarawa states of Nigeria.     

Babesia divergens becoming extinct in cattle of Northeast Hungary: new data on the past and present situation

Previously unpublished data from 1958 to 1967 attest the occurrence of Babesia divergens in cattle in several endemic foci of Northeast Hungary. During that period the number of clinical cases showed fluctuation with intervals of 4-5 years and monophasic seasonality (peaking in June). In order to assess the current status of bovine babesiosis in that region, blood samples were collected from 654 cattle on 44 farms of 36 settlements in or near the endemic area during 2005, and serum levels of IgG antibodies to B. divergens were measured by indirect fluorescent antibody test (IFAT). Only 2 samples (0.3%) showed positivity. In one village clinical babesiosis was observed over the past few years. Animals brought into the endemic area during the spring developed haemoglobinuria in the summer of the same year, but those introduced during the summer or autumn showed clinical signs only after two years. Sampled animals born and raised locally had neither haemoglobinuria nor seroconversion. Reduction in the number of cases during the past decades may have been influenced by the availability of hosts (i.e. decrease of cattle breeding) and the activity of vectors associated with climate-related changes (e.g. increase of annual sunlight hours in the endemic area). This is the first report on the prevalence of antibodies to B. divergens in cattle in Hungary.     

Risk factors for tick attachment to smallholder dairy cattle in Tanzania

A cross-sectional study was conducted in Tanga and Iringa regions of Tanzania, and a longitudinal study in Tanga, to investigate tick-control methods and other factors influencing tick attachment to the cattle of smallholder dairy farms. Most farmers reported applying acaricides at intervals of 1-2 weeks, most used acaricides that require on-farm dilution and most farmers incorrectly diluted the acaricides. Rhipicephalus appendiculatus and Boophilus spp. ticks were those most-frequently encountered on the cattle, but few cattle carried ticks of any species (only 13 and 4.6% of tick counts of the cattle yielded adult R. appendiculatus and Boophilus spp., respectively). Animals were more likely to carry one or more adult Boophilus spp. ticks if they also carried one or more R. appendiculatus adults (OR=14.4, CI=9.2, 22.5). The use of pour-on acaricides was associated with lower odds that animals carried a R. appendiculatus tick (OR=0.29, CI=0.18, 0.49) but higher odds that they carried a Boophilus spp. tick (OR=2.48, CI=1.55, 3.97). Animals >4 months old and those with a recent history of grazing had higher odds of carrying either a R. appendiculatus (ORs=3.41 and 2.58, CIs=2.34, 4.98 and 1.80, 3.71), or a Boophilus spp. tick (ORs=5.70 and 2.18, CIs=2.34, 4.98 and 1.49. 3.25), but zero-grazing management did not prevent ticks attaching to cattle even when combined with high-frequency acaricide treatments. The odds that animals carried ticks varied amongst the agro-ecological zones (AEZs) and administrative districts where the farms were situated-but there was still considerable residual variation in tick infestation at the farm level.     

The Effect of Short-interval Deltamethrin Applications to Control Tsetse on the Seroprevalence of Babesiosis in Cattle

For the past decade, treatment of cattle with 0.00375% deltamethrin (Decatix, Coopers) at two-weekly intervals has been part of an integrated approach to counteract continuous invasion of Zimbabwe by tsetse from the Mozambique fly-belt. To determine the effect of these regular deltamethrin treatments on the epidemiology of babesiosis, a survey was conducted to estimate the prevalence of antibodies against Babesia bigemina in adult communal cattle. The seroprevalence of antibodies against B. bigemina in adjacent areas, where cattle are treated with short-residual acaricides, was also determined for comparison. The prevalence of antibodies to B. bigemina was much higher in areas where dipping with a non-pyrethroid acaricide was conducted. This was attributed to the successful control of Boophilus spp. and hence a very low level of B. bigemina transmission in the 'deltamethrin treatment zone'. This low level of disease transmission was confirmed by the low prevalence of antibodies against B. bigemina in sentinel cattle that were introduced to the 'deltamethrin treatment zone'. The potential adverse effects of severely reducing the tick population should be taken into consideration at the onset of tsetse control operations in which cattle are to be treated with deltamethrin at short treatment intervals.     

Development of a novel self-medicating applicator for control of internal and external parasites of wild and domestic animals

Four trials, three in the United States and one in South Africa, were conducted to evaluate the potential value of a novel self-medicating applicator in the passive control of gastrointestinal nematodes in cattle and deer, and of files and ticks on cattle using oil-based treatments. The results of the trials demonstrated that this applicator is an effective and practical device for the passive treatment of both deer and cattle for trichostrongyle infections using the endectocide, moxidectin (Cydectin. Fort Dodge Animal Health, USA), of cattle for hom fly (Haemotobia irritans) infestations using the insecticide, cyfluthrin (CyLence, Bayer AG, Germany) and of cattle for tick infestations (in particular Amblyomma hebraeum and Rhipicephalus appendiculatus) using the acaricides deltamethrin and amitraz (Delete All, Intervet, South Africa).     

A ten-year review of commercial vaccine performance for control of tick infestations on cattle

Ticks are important ectoparasites of domestic and wild animals, and tick infestations economically impact cattle production worldwide. Control of cattle tick infestations has been primarily by application of acaricides which has resulted in selection of resistant ticks and environmental pollution. Herein we discuss data from tick vaccine application in Australia, Cuba, Mexico and other Latin American countries. Commercial tick vaccines for cattle based on the Boophilus microplus Bm86 gut antigen have proven to be a feasible tick control method that offers a cost-effective, environmentally friendly alternative to the use of acaricides. Commercial tick vaccines reduced tick infestations on cattle and the intensity of acaricide usage, as well as increasing animal production and reducing transmission of some tick-borne pathogens. Although commercialization of tick vaccines has been difficult owing to previous constraints of antigen discovery, the expense of testing vaccines in cattle, and company restructuring, the success of these vaccines over the past decade has clearly demonstrated their potential as an improved method of tick control for cattle. Development of improved vaccines in the future will be greatly enhanced by new and efficient molecular technologies for antigen discovery and the urgent need for a tick control method to reduce or replace the use of acaricides, especially in regions where extensive tick resistance has occurred.     

Productivity and health effects of anaplasmosis and babesiosis on Bos indicus cattle and their crosses, and the effects of differing intensity of tick control in Australia

Tick fever is an important disease of cattle where Rhipicephalus (Boophilus) microplus acts as a vector for the three causal organisms Babesia bovis, Babesia bigemina and Anaplasma marginale. Bos indicus cattle and their crosses are more resistant to the clinical effects of infection with B. bovis and B. bigemina than are Bos taurus cattle. Resistance is not complete, however, and herds of B. indicus-cross cattle are still at risk of babesiosis in environments where exposure to B. bovis is light in most years but occasionally high. The susceptibility of B. indicus cattle and their crosses to infection with A. marginale is similar to that of B. taurus cattle. In herds of B. indicus cattle and their crosses the infection rate of Babesia spp. and A. marginale is lowered because fewer ticks are likely to attach per day due to reduced numbers of ticks in the field (long-term effect on population, arising from high host resistance) and because a smaller proportion of ticks that do develop to feed on infected cattle will in turn be infected (due to lower parasitaemia). As a consequence, herds of B. indicus cattle are less likely than herds of B. taurus cattle to have high levels of population immunity to babesiosis or anaplasmosis. The effects of acaricide application on the probability of clinical disease due to anaplasmosis and babesiosis are unpredictable and dependent on the prevalence of infection in ticks and in cattle at the time of application. Attempting to manipulate population immunity through the toleration of specific threshold numbers of ticks with the aim of controlling tick fever is not reliable and the justification for acaricide application should be for the control of ticks rather than for tick fever. Vaccination of B. indicus cattle and their crosses is advisable in all areas where ticks exist, although vaccination against B. bigemina is probably not essential in pure B. indicus animals.     

Cattle farmers' perception of the nuisances caused by insects and ticks to cattle in the Canton Jura

During summer 2005, a survey of 172 farmers from the Canton Jura was carried out to determine their perception of the nuisances caused by insects and ticks to cattle. The presence of ticks was significantly more often spontaneously mentioned by farmers in the Clos-du-Doubs and in the district of Delémont (95% and 72%, respectively) than in Ajoie and in the Franches-Montagnes (29% and 19%, respectively, p < 0.0001). 20% of the farmers perceived the tick populations to be increasing. The following diseases were spontaneously listed: keratoconjunctivitis (59%), mastitis related to flies (31%), hypodermosis (15%), ehrlichiosis (12%) and babesiosis (10%). Suspicion of ehrlichiosis was significantly more often mentioned in the district of Delémont than in the rest of the canton (p < 0.001) which was associated with the presence of ticks observed by farmers (p < 0.001). Cases of clinical babesiosis, which is common in the Clos_du_Doubs, were mentioned by farmers outside this area: Delémont (3/57) and Ajoie (1/26). In this study babesiosis depends on the production area, on the presence of ticks observed by farmers and on the presence of water in the pasture (p < 0.05). This study shows that cattle farmers are well aware of the presence of the diseases linked to insects and ticks and that the perception of their distribution is in accordance with the epidemiological data. An overview of the epidemiology of tick- and insect-related diseases can therefore be established by surveying farmers' perception, but should then be confirmed by a biomedical study. Due to their accurate observations, farmers are key participants at the first level of surveillance systems of animal diseases.     

Factors related to cattle infestation level and resistance to acaricides in Boophilus microplus tick populations in New Caledonia

Boophilus microplus, even in the absence of babesiosis, is a major disease of the cattle in New Caledonia where the particular farming system associates continental European breeds and a tropical climate tempered by the Pacific Ocean. In order to have a better understanding of the factors involved in cattle tick infestation, to decrease the possible wastage and use of chemicals and to increase the lifetime of the acaricides, the veterinary authorities investigated the conditions of the chemical treatments. A survey among 148 cattle farms of the whole of New Caledonia was carried out and factors that explain the development of tick resistance and cattle infestation have been determined. From this survey, three typologies for the main characteristics of the farms have been set up, the technical practices of the farmers and the tick control measures applied by the farmers, respectively. Some variables are significantly associated with the tick resistance to deltamethrin but their contribution to the explanation model is always moderate: farms in the south, with a positive resistance gradient from east to west, absence of bush fire and membership to a cattle farmers organization. The more the farmers have intensified their breeding-male castration, weaning, heifer separation, drenching, etc.-and pasture-high stock rate, mowing, extra feeding of the cows, many paddocks, etc.-techniques, the higher was the probability for the ticks in their farm to be resistant to deltamethrin. The technical details of the acaricide treatment had a low contribution to the explanation model. However, the use of a spray generated more resistance than a dip. Furthermore, there is a negative resistance gradient when the farmers increased the treatment interval average. Considering infestation, none of the variables from the three typologies were associated with the two infestation variables (1: semi-engorged tick females and 2: other ticks) at the herd level. However, the seven studied variables-the three typologies, breed, age, body condition score and breeding status-affected significantly the two infestation variables at the cow level, but their predictive ability remained very low (R(2)<3.5%). This result-individual effect more important than herd effect on the infestation-is confirmed by the importance of the variance of the intra-farm factors (99%) when compared with inter-farm factors (1%). Cows of Charolais breed, in poor body condition, old, pregnant or lactating, and those of the farms with irrational and high pressure control of ticks are the most infested.     

Differential Bos taurus cattle response to Babesia bovis infection

Bovine babesiosis is a tick-borne disease caused by Babesia spp. haemoprotozoans. The disease is of great importance at tick enzootic unstable areas and hampers cattle production in several developing countries. The available immunisation alternatives are pre-immunition and attenuated vaccines. Despite being efficient and protective, they are unsafe as they use cattle blood cells as inoculum and may potentially spread other diseases. Another alternative to help in babesiosis control would be the identification of genetically resistant cattle to Babesia bovis infection. The objective of this work was to phenotype cattle based on primary response against B. bovis infection. Two-hundred and forty half-sib Hereford and Aberdeen Angus heifers (120 animals from each breed), 12-18-month-old na?ve cattle, originated from a tick-free area in Southern Brazil, were used in the experiment. Animals were monitored following an inoculation with 1x10(7)B. bovis parasitised erythrocytes. Results showed three different phenotypes: 1-'susceptible', animals with babesiosis clinical signs that received treatment to avoid death; 2-'intermediate', animals with clinical signs: parasitaemia, >or=21.5% reduction in packed cell volume (PCV) and increase in body temperature when compared to their pre-challenge physiological parameters, no specific treatment was needed as animals self recovered from the disease, and 3-'resistant', animals without clinical signs that showed B. bovis presence in blood smears, <21.5% PCV reductions, with little or no increase in body temperature and no need for babesiosis treatment. The frequencies of each phenotype were: 45.4, 26.7, and 27.9%, respectively, demonstrating the existence of phenotypic variation for B. bovis in Bos taurus cattle.     

Inhibition of Babesia divergens in cattle by oxytetracycline

The effects of continuous oxytetracycline administration on the development of parasitaemia of Babesia divergens during both natural and artificial infections were studied. During natural exposure on grazing heavily infested with Ixodes ricinus, seven out of 42 cattle with no previous exposure to tick-borne diseases were injected every four days with a long acting preparation of oxytetracycline at a dose rate of 20 mg/kg. During the six week grazing period 21 untreated cattle developed a patent parasitaemia of B divergens and all became seropositive by the fluorescent antibody test. In contrast, no parasites were observed in treated cattle and antibody titres remained low. Artificial infections were studied with different dose levels of oxytetracycline and their effects on antibody stimulation noted. First, four groups of cows were infected with 10(8) erythrocytes infected with B divergens, three groups being injected every four days with the long acting oxytetracycline formulation at dose levels of 20, 10 and 5 mg/kg, respectively. The highest level completely inhibited parasite replication and antibody formation; the same was observed in one animal dosed at 10 mg/kg but the remainder, plus those treated at 5 mg/kg, developed both low parasitaemia and high antibody titres. The untreated cows developed severe babesiosis. A further untreated control group was added and three weeks after cessation of oxytetracycline treatment all were infected with 10(9) erythrocytes infected with a homologous isolate of B divergens. The controls, plus those in which the previous infection had been completely inhibited, developed severe clinical babesiosis but the remainder were refractory to parasite development.     

Anaplasma infection in free-ranging Iberian red deer in the region of Castilla-La Mancha, Spain

Organisms in the genus Anaplasma are obligate intracellular pathogens that multiply in both vertebrate and invertebrate hosts. The type species, Anaplasma marginale, causes bovine anaplasmosis and infects erythrocytes of the vertebrate host and undergoes a complex developmental cycle in ticks which serve as biological vectors. Infected cattle, wild ruminants and ticks can all serve as reservoirs of A. marginale. In this study, hunter killed Iberian red deer (Cervus elaphus hispanicus) from the region of Castilla-La Mancha in southwestern Spain were tested for Anaplasma infection. We found that 10% of the deer examined were seropositive for Anaplasma. Three A. marginale strains were subsequently obtained from salivary glands of Hyalomma marginatum that were removed from these deer, and the sequence of the major surface protein (msp)4 gene was determined for each strain and used for phylogenetic studies. Maximum parsimony analyses of msp4 sequences from H. marginatum ticks in comparison with New World cattle and bison isolates reported previously, suggested different origins for these Spanish A. marginale strains. The results of this study demonstrated that Iberian red deer are naturally infected with Anaplasma, and may therefore serve as a wildlife reservoir of the pathogen. Although the link between deer infection and the strains of A. marginale identified in ticks was not established, H. marginatum and Rhipicephalus bursa were identified as potential biological vectors for A. marginale in this region and may effect transmission of A. marginale between deer and cattle populations.     

A Vegetation Index qualifying pasture edges is related to Ixodes ricinus density and to Babesia divergens seroprevalence in dairy cattle herds

Babesia divergens, transmitted by the tick Ixodes ricinus, is the main agent of bovine piroplasmosis in France. This Apicomplexa often is present in asymptomatic carriers; however, clinical cases are rare. While numerous factors are known to influence tick density, no risk factor of contact with B. divergens has been identified for cattle. Our study aimed to explore whether a Vegetation Index could serve as an indirect indicator of within-herd B. divergens seroprevalence. In February 2007, blood samples were taken from all of the cows in 19 dairy cattle herds in Western France and IFAT serology was performed individually to measure B. divergens seroprevalence. The following spring, I. ricinus nymphs were collected by drag sampling along transects on the vegetation of each farm's pasture perimeters. Tick density was related significantly to a Vegetation Index (V.I., ranging from 1 to 5) that took into account the abundance of trees and bushes on the edge of pastures: most ticks (57%) were found in transects with the highest V.I. (covering 15% of the explored surface in the study area). At the farm level, the proportion of transects presenting I. ricinus nymphs was significantly related to B. divergens seroprevalence: the farms with more than 15% of transects with I. ricinus had a significantly higher risk of high seroprevalence. The proportion of pasture perimeters where the V.I.=5 also was significantly related to B. divergens seroprevalence: the farms where more than 20% of transects had a V.I.=5 had a significantly higher risk of high seroprevalence. Given that the Vegetation Index is a steady indicator of the potential I. ricinus density in the biotope, we recommend that the risk of high B. divergens seroprevalence in cows be evaluated using this tool rather than drag samplings.     

Evaluation of historical factors influencing the occurrence and distribution of Mycobacterium bovis infection among wildlife in Michigan

OBJECTIVES: To determine historical events leading to establishment of bovine tuberculosis in the white-tailed deer population in the northeastern corner of the lower peninsula (NELP) of Michigan and describe factors relevant to the present outbreak of bovine tuberculosis in Michigan. SAMPLE POPULATION: Cattle and white-tailed deer in Michigan from 1920 to 1990. PROCEDURES: A search of extant historical documents (eg, scientific journals, books, public reports, and correspondence and internal reports from governmental agencies) was conducted. Factors investigated included the number of cattle and prevalence of tuberculosis, deer population and density levels, and changes in regional environments affecting the population and management of cattle and wild deer. RESULTS: High deer numbers and severe winter feed shortages resulting from habitat destruction in the NELP in 1930 contributed to the transmission of tuberculosis from cattle to deer. Starvation increased the susceptibility of deer to infection and modified behavior such that exposure to infected cattle was increased. Relocation of deer resulted in spread of infection to other sites, including locations at which spatial clusters of tuberculosis presently exist. Ribotyping of Mycobacterium bovis from a human patient suggests that the strain of M. bovis presently infecting white-tailed deer in the region is the same strain that affected cattle farms at that time. CONCLUSIONS AND CLINICAL RELEVANCE: Feeding deer to maintain numbers above the normal carrying capacity of the NELP led to deer depending on consumption of livestock feed for survival during winter and increased contact with domestic cattle. This practice should be avoided.