Canine angiostrongylosis: the French heartworm: an emerging threat in North America

Angiostrongylus vasorum, French heartworm, is a metastrongloid parasite found in the pulmonary arteries and right ventricle of wild and domestic canids and various other animals. The natural definitive hosts are species of foxes. The geographic distribution of the parasite includes various countries of Europe, Africa, South America, and North America. Angiostrongylosis is considered an emerging disease in dogs in Europe. In North America, autochthonous A. vasorum infection occurs only in the Canadian province of Newfoundland-Labrador. Computer modeling suggests there is a high probability that A. vasorum will spread to other parts of North America and will likely become endemic in the eastern half of the continent and in the states and provinces along the western coast. Animals acquire infection by the ingestion of gastropod or frog intermediate hosts that carry the infective 3rd-stage larvae. Frogs can also serve as paratenic hosts. Definitive antemortem diagnosis is by detection of L(1) in feces, sputum, or bronchoalveolar lavage samples. Baermann fecal examination is the most reliable method for fecal detection. However, false negative results can occur due to the typical erratic/sporadic fecal larval shedding pattern of A. vasorum. Recently, promising new methods for A. vasorum infection diagnosis have been reported involving polymerase chain reaction of blood and fecal samples and a sandwich ELISA for detection of circulating worm excretory/secretory antigen. Current treatment options include moxidectin, milbemycin oxime, and fenbendazole.     

Autochtonous infection of dogs and slugs with Angiostrongylus vasorum in Hungary

On the course of a helminthological survey of the dogs of Baranya County, Hungary Angiostrongylus vasorum infection was detected in two asymptomatic dogs. Identification of the parasite was based on morphology of the first-stage larvae (L1) isolated from droppings, and successful experimental infection with first stage larvae to laboratory reared Discus rotundatus and Lissachatina fulica snails, in order to exclude species of the family Filaroididae that have similar larvae to A. vasorum. While angiostrongylosis is widespread among foxes, this is the first report of A. vasorum infection in housedog in Hungary. In gardens, where infected dogs were being kept 91 specimens of 6 species of limacid and arionid slugs were collected of which 5 specimens of Arion lusitanicus were found to carry larvae of A. vasorum. Dogs usually do not ingest such large slugs willingly. Frogs are known to act as paratenic hosts in the life cycle of A. vasorum. Since one of the infected dogs harboured also infection with the intestinal trematode Alaria alata, of which frogs certainly play the role of the second intermediate host, therefore it is assumed that in this case the dog became infected with A. vasorum by eating frogs.     

Autochthonous infections with Angiostrongylus vasorum in dogs in Switzerland and Germany

Angiostrongylus vasorum is endemic in foxes an other carnivores in the South and south East of France, Denmark and Great Britain. The reddish nematode is present in the Arteria pulmonalis and the right side of the heart and causes respiratory and cardiovascular symptoms. From 1999 to 2004, A. vasorum was diagnosed in 5 dogs from northern Switzerland, in 1 dog from southern Germany and in 3 dogs from south Ticino. Clinical signs in the affected dogs varied and ranged from cough, tachypnoea and dyspnoea to neurological symptoms in 2 of the dogs. Four dogs died and in 3 of the 4 dogs adults and larvae were found after digestion of formalin-fixed tissue. Diagnosis in the other 5 dogs was achieved by detecting the larvae coproscopically. Based on anamnestic data, these 9 dogs are probably autochthonous cases.     

On pseudorabies in carnivores in Denmark. II. The blue fox (Alopex lagopus)

In the autumn of 1968 Pseudorabies was demonstrated in blue foxes (Alopex lagopus) on two fur farms in Jutland. The foxes had been fed with dead piglets from herds of swine belonging to the two farmers.At the same time the disease was diagnosed in piglets from these herds. At the two fur farms there were, respectively, 34 and eight foxes in all; respectively 26 and two died with signs of pruritus. Only few of the surviving foxes had had the opportunity to eat piglets. Eight weeks after the period of disease no antibodies against psedorabies virus could be demonstrated in serum of three surviving foxes.The virus content in various organs of six foxes submitted to the Laboratory was determined by titration in tissue cultures (Table 1).The results indicate that the spreading of virus from the port of entry of the virus to the brain must have occurred by nervous routes.The presence of virus in the tonsils of five of the six foxes and in the nasal mucosa of the sixth one, together with similar results from examination of dogs at this institute would seem to suggest that a transmission of the disease from these carnivores to other animals is possible.Based upon the high virus titers in the medulla oblongata and pons, and with reference to similar results of examination of dogs and to a previous publication on Pseudorabies in Danish red foxes, it is concluded that these tissues must be the material of choice for demonstration of Pseudorabies virus in these carnivores.     

Serologic detection of Angiostrongylus vasorum infection in dogs

Angiostrongylus vasorum, French Heartworm, is a metastrongylid nematode infecting the pulmonary arteries and right heart of wild and domestic canids in various regions of the world. Infection in dogs can result in fatal cardiopulmonary disease. A single endemic focus of A. vasorum in North America occurs in the southeastern portion of Newfoundland, Canada. Dogs are currently diagnosed by detection of first-stage larvae shed in feces using the Baermann technique or fecal flotation. However, these procedures may lack sensitivity due to intermittent fecal larval shedding. The potential for using detection of circulating worm antigen for diagnosis was investigated by developing a sandwich-ELISA using rabbit anti-whole adult worm antiserum. This test detected circulating antigen in sera from 22/24 Baermann positive dogs naturally infected with A. vasorum. Negative results (0/52) were obtained from sera collected from Baermann negative dogs from outside of the endemic region, and from sera (0/30) from dogs from non-endemic regions that were infected with Crenosoma vulpis, the fox lung worm. Receiver operating curve analysis gave a specificity of 100% and a sensitivity of 92% for the sandwich-ELISA at an optical density cut-off of 0.19. Subsequently, 239 dogs from Newfoundland displaying clinical signs of cardiopulmonary disease, were examined using both the Baermann fecal examination and the sandwich-ELISA. Larvae were detected in 10% (24/239) of these dogs by fecal examination, whereas the sandwich-ELISA detected circulating antigen of A. vasorum in serum from 18.8% (45/239) of the dogs. This suggests that fecal diagnostics may have missed approximately half of the A. vasorum infected dogs, and that the sandwich-ELISA may be a useful tool in the diagnosis of this parasite.     

An epidemic of sylvatic rabies in Finland--descriptive epidemiology and results of oral vaccination.

When rabies reappeared in Finland in April 1988, the country had been rabies free since 1959. Soon a picture of sylvatic rabies become evident, its main vector and victim being the raccoon dog (Nyctereutes procyonoides). Between 8 April 1988 and 16 February 1989, 66 virologically verified cases were recorded (48 raccoon dogs, 12 red foxes, 2 badgers, 2 cats, 1 dog and 1 dairy bull) in an area estimated at 1700 km2 in south-eastern Finland. The greatest distance between recorded cases was 67 km. A positive reaction with monoclonal antibody p-41 indicated that the virus was an arctic-type strain. A field trial on oral immunization of small predators was initiated in September 1988 using Tübingen fox baits according to the Bavarian model of bait distribution. Each bait contained 5*10(7) TCID50/ml modified live rabies virus (SAD-B19). The 6 months' surveillance indicate a seroconversion rate of 72% (N = 126) in the raccoon dog population, 67% (N = 56) in the red foxes and 13% (N = 16) in the badgers, when titers greater than or equal to 1.0 IU/ml are considered seropositive. In the whole follow-up period, no statistically significant difference could be detected between the raccoon dogs and red foxes in the rate of seroconversion or in the uptake of tetracycline from the baits. Notably high antibody levels were recorded in both raccoon dogs and red foxes within 4-5 months after vaccination. Of the seropositive animals, the proportion of animals with titers 3.0 IU/ml or greater was higher in raccoon dogs (73%) than in red foxes (51%) (x2 = 5.29, p less than 0.05). The trial shows that raccoon dogs can be immunized against rabies in the field with vaccine baits originally developed for controlling sylvatic rabies in foxes.     

Urban domestic dog populations as a source of canine distemper virus for wild carnivores in the Coquimbo region of Chile

Urban areas can support dog populations dense enough to maintain canine distemper virus (CDV) and can be a source of infection for rural dogs and free-ranging carnivores. The aim of this study was to investigate the relationships between urban and rural domestic dog and wild carnivore populations and their effects on the epidemiology of CDV to explain retrospectively a CD outbreak in wild foxes in 2003. From 2005 to 2007 a cross-sectional household questionnaire survey was conducted in Coquimbo and Ovalle cities, in three towns and in rural sites along two transects from these cities to the Fray Jorge National Park (FJNP) in the Coquimbo region, Chile. Blood samples were collected from unvaccinated dogs at surveyed households and from free-ranging foxes in rural areas along the transects. The seroprevalence of CDV in domestic dogs was higher in urban than in rural areas and in the later was highest in dogs born before 2001-2002. The seroprevalence of CDV in foxes was higher in areas closer to human settlements. A high seroprevalence in dogs born before 2001-2002 further supports a link between CDV patterns in rural dog and fox populations. In our study area, urban dogs are proposed to be the source of CDV infection to wild carnivores. The large dog population size and density detected in Coquimbo and Ovalle provides optimal conditions for maintaining a large and dense susceptible population of dogs, which can act as a reservoir for highly infectious diseases and could have been the source of infection in the CD outbreak in wild foxes.     

Spirocerca lupi isolated from gastric lesions in foxes (Vulpes vulpes) in Sicily (Italy)

Spirocerca lupi (Rudolphi 1809) is a cosmopolitan nematode of dogs and wild carnivores. In the past it has been reported in Italy, mainly in southern regions and in Sicily, where the parasite was observed in foxes in 2005. The parasite typically produces nodular masses in the oesophagus and thoracic aorta. During the 2003-2004 hunting season, the authors investigated a total of 55 foxes (Vulpes vulpes) hunted or killed by car accidents in the provinces of Palermo and Agrigento. All the foxes were subjected to necropsy and 6 (9.16%) had S. lupi nodules located exclusively in the gastric wall. The nature of the nodules was determined by opening them and detecting the nematodes inside, which were identified as S. lupi. Some of the nodules were characterized anatomopathologically and histopathologically. The formation of the parasitic nodule in the stomach only suggests a deviation from the route commonly followed by the nematode to reach the oesophagus, the elective anatomical site for completion of its lifecycle. This survey gives a contribution to the epidemiology of this parasite which is severely outdated in Italy and highlights some distinctive features of the life cycle and parasite migration.     

Characterization of widespread canine leishmaniasis among wild carnivores from Spain

Visceral Leishmaniasis (VL) is an emerging zoonotic parasitic disease caused by Leishmania infantum in Mediterranean countries, with sand flies (Phlebotomus spp.) as vectors and dogs as the main domestic reservoir. The role of wild carnivores in the epidemiology of leishmaniasis is still controversial. In order to determine the prevalence of natural infection with L. infantum in wild carnivores from Spain, we analyzed 217 samples by PCR and western blotting and used restriction fragment length polymorphism (RFLP) to compare the patterns present in wild carnivores with those of domestic dogs from the same areas. DNA of the parasite was detected in spleen or blood samples from 35 (16.12%) analyzed wild carnivores, including 8 of 39 (20.5%) wolves (Canis lupus), 23 of 162 (14.1%) foxes (Vulpes vulpes), 2 of 7 (28.6%) Egyptian mongooses (Herpestes ichneumon), 1 of 4 genets (Geneta geneta), and 1 of 4 Iberian lynxes (Lynx pardinus). No significant sex or age differences in prevalence were observed in wolves and foxes (P>0.05), but there was a significant difference among regions in foxes (P<0.05). A total of 12 PCR-RFLP patterns were found in foxes, 6 in wolves, 4 in dogs, 2 in Egyptian mongooses and 1 in lynx and genet. RFLP patterns differed between dogs and foxes in the two areas where they could be compared. This is the first study of canine leishmaniasis in wild canids and other carnivores from different regions of Spain by PCR. The prevalence of infection indicates the existence of natural infection in apparently healthy wild carnivore populations, and our results are suggestive of a sylvatic cycle independent of dogs.     

Angiostrongylus vasorum and Eucoleus aerophilus in foxes (Vulpes vulpes) in Great Britain

The nematode parasite Angiostrongylus vasorum is a source of increasing concern in several parts of the world, where it causes significant disease in dogs. Wild canids, especially foxes, are likely to have a role in the epidemiology of canine infection, and the parasite could also affect fox health and population dynamics. The heart and pulmonary vasculature of 546 foxes culled mostly by gamekeepers in Great Britain in 2005–2006 were examined by dissection and a modified flushing technique. Forty foxes were found to be infected, giving an overall prevalence in the UK fox population of 7.3% (5.3–9.9). Prevalence varied widely between regions, from 0% (0–3) in Scotland and northern England to 23% (16–32) in south-east England. This closely matches the perceived incidence of disease in dogs, which is commonly diagnosed in the south-east but rarely in the north. In the Midlands, where disease has recently appeared in dogs, prevalence in foxes was 4.8% (2–11). Close geographical overlap of parasite distribution in foxes and dogs does not necessarily indicate an important wildlife reservoir of infection, but does suggest that A. vasorum might be spreading northwards. The hearts of infected foxes had thicker right ventricles than those of uninfected foxes, suggesting that the parasite could affect fox health and fitness. Burdens ranged from 1 to 59 adult nematodes. Sex, age and body condition were not significantly associated with infection. Eucoleus aerophilus and Crenosoma vulpis, nematode parasites of the respiratory system, were found in 213 and 11 foxes respectively, with slightly higher prevalence of E. aerophilus in the south and east. No specimens of the heartworm Dirofilaria immitis were found, giving an upper 95% confidence interval for prevalence of 0.84%.     

Canine angiostrongylosis: an emerging disease in Europe

Objective – The aim of this article is to review Angiostrongylus vasorum infection in dogs, including the life cycle, signalment, clinical signs, diagnosis, and treatment. Apparent changes in the epidemiology of this unique parasite are considered, alongside information available regarding its recent geographic spread.
Etiology – A. vasorum is a metastrongyloid parasite capable of causing an array of clinical problems in dogs, including cardiorespiratory, coagulopathic, and neurologic signs. Currently, the parasite has a worldwide distribution; however, it usually arises in small pockets of enzootic foci. Recent reports suggest a changing distribution of this parasite, which has renewed interest in its epidemiology and in the risk of expansion to new areas including mainland North America.
Diagnosis – A definitive diagnosis of angiostrongylosis is usually made using the modified Baermann technique either using feces or tracheobronchial secretions; however, this review also discusses novel methods such as serologic and molecular techniques.
Therapy – Once a diagnosis of angiostrongylosis is made, prompt treatment should follow with anthelmintic drugs (such as moxidectin/imidacloprid, milbemycin oxime, or fenbendazole) and supportive care dependent upon the patient's clinical signs. Currently, there is no proven prophylactic regime.
Prognosis – The prognosis appears to be very dependent upon the severity of clinical signs at presentation. A. vasorum can be fatal and death may be sudden. However, if a prompt diagnosis is made and appropriate treatment is administered complete clinical resolution is possible.     

Acute haemoabdomen associated with Angiostrongylus vasoruminfection in a dog: a case report

A one-year-old intact female, Danish shorthaired pointer was referred to the emergency service with a history of acute collapse and pale mucous membranes after a month of reduced activity but with no other clinical signs. An ultrasound examination of the abdomen indicated the presence of a large amount of free fluid with no obvious cause such as neoplasia or splenic rupture. Fluid analysis had the macroscopic appearance of blood with no signs of infection or neoplasia. Multiple Angiostrongylus vasorum L1 larvae were revealed on a direct rectal faecal smear. The dog was treated with fenbendazole 25 mg/kg orally once daily for 20 days and given supportive treatment. The dog was stabilised on this treatment. Haemoabdomen is a clinical sign where surgical intervention is often considered an integral part of the diagnostic investigation (i.e., laparotomy) or treatment. Failing to make the diagnosis of canine angiostrongylosis before performing surgery may have a serious adverse affect on the outcome. Consequently, in areas where A. vasorum is enzootic, a Baermann test and a direct faecal smear should be included in the initial diagnostic investigation of all dogs presenting with bleeding disorders of unknown origin.     

Prevalence of Dirofilaria immitis in a stray dog population in western New York.

In a survey in 1973 undertaken to determine the prevalence of Dirofilaria immitis in 100 dogs from a public pound in western New York, 2 dogs were found to be positive. Microfilariae of Dipetalonema sp were not present. Hearts and pulmonary blood vessels were dissected for adult worms, and blood samples were examined for microfilariae, using a modified Knott technique. Although adult heartworms and microfilariae have been recovered from wild foxes, coyotes, and wolves in New York, this is the first report of heartworm in domesticated dogs in the state.     

Studies on the role of the red fox (Vulpes vulpes) as a potential definitive host of Neospora caninum

Neospora (N.) caninum is a protozoan parasite which is regarded as a major cause of abortion in cattle. Dogs and coyotes are definitive hosts of N. caninum which may shed environmentally resistant stages, oocysts, in their feces. Epidemiological studies in Germany showed that the presence of dogs increased the risk of a bovine herd to be N. caninum-positive in a bulk-milk ELISA test. However, there were also N. caninum-positive herds where dogs were not kept together with cattle.This leads to the question whether canids other than dogs, e.g., foxes, might be involved in the horizontal transmission of N. caninum. Therefore, the aim of our examinations in wild animals was to find out whether there are indications for a sylvatic cycle with foxes as definitive hosts and deer, roe deer and wild mice samples contained structures which resembled those of coccidian oocysts. In 13 of these 65 samples coccidian DNA was detected using a 18S rRNA gene based polymerase chain reaction (PCR).The examination of the 65 samples in a N. caninum-specific PCR revealed no positive result. Hammondia (H.) heydorni-DNA was detected in two samples. In addition, brain samples from 528 foxes, 224 wild mice, 16 deer and roe deer as well as from 1 wild boar were examined for the presence of N. caninum DNA by real time PCR. All samples tested negative by PCR. In conclusion, our study yielded no evidence indicating that the examined animals were part of a sylvatic cycle for N. caninum.     

Intracranial haemorrhage associated with Angiostrongylus vasorum infection in three dogs

This report describes three dogs with intracranial haemorrhage secondary to severe coagulation defects associated with Angiostrongylus vasorum infection. The initial case was diagnosed at necropsy, with two subsequent cases diagnosed antemortem and successfully treated. The dogs ranged in age from 14 months to four years and were presented for evaluation of a severe, subacute onset of suspected cerebral disease. Magnetic resonance imaging performed on all three dogs was suggestive of multiple areas of intraparenchymal brain haemorrhage. Coagulation assays showed a consumptive coagulopathy resembling chronic disseminated intravascular coagulation. Postmortem examination of the initial case confirmed the presence of multiple intracranial and extracranial haemorrhages. An unexpected finding was that of a marked multifocal nematode infection of the lungs with an associated vasculopathy. The parasites were confirmed to be A vasorum. In the two other dogs, faecal examination by Baermann technique confirmed A vasorum infection. Both dogs were treated with fenbendazole and one was additionally given a plasma transfusion. Repeated coagulation assays were normal within one week. Neurological examinations were normal for both dogs within six weeks. This case series indicates that A vasorum infection should be considered as a possible aetiology of intracranial haemorrhage in dogs.     

The so-called "fox tapeworm"

The so-called "fox tapeworm" (Echinococcus multilocularis), the causative agent of a severe disease in man (alveolar echinococcosis), is presently under public discussion in Switzerland. Therefore, actual information is provided on the life cycle of the parasite, epidemiology, disease in humans, symptomatology, diagnosis, therapy and prophylaxis. It is recommended that in endemic regions hunters handling foxes should wear protective gloves, dead foxes should be transported in plastic bags and wild fruits, berries and vegetables should be carefully washed and--if possible--heated to more than 70 degrees C for some minutes prior to consumption. After contact with foxes or other final hosts (dogs, cats) infected with E. multilocularis, persons should be monitored with the highly sensitive and specific Em2-ELISA for serum antibodies aiming at an early diagnosis and treatment of a potential infection.     

Veterinary aspects of alveolar echinococcosis--a zoonosis of public health significance

Human alveolar echinococcosis (AE), caused by the metacestode stage of Echinococcus multilocularis, is a serious zoonosis which caused up to 100% lethality in untreated patients before the 1970s, when modern methods of treatment were not yet established. AE occurs in large areas of the northern hemisphere mostly with low country-wide prevalences, but high prevalences of up to 4% have been reported from small population groups in highly endemic foci, e.g. from China. AE includes many veterinary aspects which are the topic of this review. Recent studies have shown that E. multilocularis has a wider geographic range than previously anticipated. There is evidence for growing populations of red foxes (Vulpes vulpes) in some areas, for increasing invasion of cities by foxes and also for establishment of the parasite cycle in urban areas. These and other factors may lead to an increased infection risk for humans. Significant progress has been made in the development of sensitive and specific new techniques for the intra vitam and post mortem diagnosis of intestinal E. multilocularis infection in definitive hosts, notably the detection of coproantigen by enzyme-linked immunosorbent assay and of copro-DNA by PCR. Both tests can also be used for the identification of E. multilocularis in faecal samples collected in the environment. Recommendations are given for chemotherapy and chemoprophylaxis of the intestinal infection in definitive hosts. In recent years, infections with the metacestode stage of E. multilocularis have not only been diagnosed in humans in several regions, including at least eight countries in central Europe, but also in animal species which do not play a role in the transmission cycle (wild and domestic pigs, dogs etc.). From 1987 to 2000 our group in Zurich has diagnosed 10 cases of AE in dogs and 15 in captive monkeys. In 2 dogs, concurrent infections of the intestine and of the liver with adult and larval stages of E. multilocularis, respectively, were observed for the first time. Clinical data are presented, and methods of diagnosis and treatment (surgery, chemotherapy) are described. Furthermore, small liver lesions caused by E. multilocularis were diagnosed in 10% of 90 slaughter pigs, and 2.9% of 522 breeding sows had specific serum antibodies against parasite antigens. In view of the unpredictable epidemiological situation, all possible measures for preventing E. multilocularis infections in humans and in domestic animals should be initiated by the veterinary and health authorities.     

Biochemical serum profiles in dogs experimentally infected with Angiostrongylus vasorum (Baillet, 1866)

The biochemical profiles of crossbred dogs experimentally infected with the parasite Angiostrongylus vasorum were studied. Two groups of five dogs were experimentally inoculated with 50 and 100 third stage infective larvae (L3) of A. vasorum per kilogram of body weight. A third group of five uninfected animals were used as control. Serum from these animals were used for biochemical tests to measure total and fractioned proteins, urea, creatinine and to determine the activities of aspartate (AST), alanine (ALT) aminotransferase, gamma-glutamyl transferase (GGT), alkaline phosphatase (PAL) and creatine kinase isoenzyme MB (CK-MB). The alpha-1, alpha-2 and beta-globulins fractions showed alterations during acute phase of the infection. No modifications were observed in the biochemical profiles of ALT, AST, GGT, PAL, urea and creatinine. CK-MB was shown to be a good early indicator of cardiac injury in dogs experimentally infected with A. vasorum.     

Long-term study of Sarcoptes scabiei infection in Norwegian red foxes (Vulpes vulpes) indicating host/parasite adaptation

The red fox (Vulpes vulpes) population, in Norway, was na?ve to Sarcoptes scabiei prior to the late 1970s when this parasite was first recorded and a still ongoing epidemic started. During the course of this protracted epidemic some degree of host/parasite adaptation, with the occurrence of healthy antibody positive foxes, might be expected. In the present study the prevalence of sarcoptic mange and serologically identified S. scabiei exposure was investigated in 363 Norwegian red foxes, shot by hunters during two different study periods (1994-1995 and 2002-2005). The sarcoptic mange diagnosis was based upon the presence of clearly visible lesions in the skin of the cadaver with confirmatory demonstration of S. scabiei. The serodiagnosis was based on an indirect-ELISA. There was a significant decrease in prevalence of both mange cases and seropositive animals from the first to the second study period. Whilst the mange prevalence fell more than threefold, from 30.0% to 6.6%, the seroprevalence dropped less dramatically from 53.3% to 19.1%. The smaller decrease in seroprevalence compared to mange cases reflected a significantly higher ratio of seropositive-mange negative versus seropositive-mange positive foxes, during the second study period, 40:18, compared to the first, 14:18. These findings indicate that the red fox population is adapting to live with the parasite and that low-grade or sub-clinical infections, and even recoveries, occur amongst exposed foxes. Mange positive foxes had significantly poorer body condition than those without sarcoptic mange. No significant difference in body condition was seen between seropositive-mange negative versus seronegative-mange negative foxes. The ELISA sensitivity was found to be 95% and proved a useful tool for investigating the exposure to S. scabiei in wild foxes. This study is believed to be the first pointing to a long-term Sarcoptes/fox adaptation, combining long-term prevalence studies of clinical sarcoptic mange and serological evidence of exposure to the parasite in the general fox population.