REVIEW

Neosporosis and hammondiosis

in dogs
The dog is a definitive host of the protozoan parasite Neospora Toxoplasma gondii infection (Dubey and
others 1988). Subsequent retrospective ana-
caninum, and in many parts of the world, infection is relatively lyses have now re-classified a number
common as determined by serology. Reported seroprevalences of cases as due to N caninum infections
(Dubey and others 1988, Munday and
usually range from 0 to 20 per cent, however, reports of clinically others 1990, Patitucci and others 1997,
affected dogs are infrequent. Affected dogs are generally less than Ellis 1998). Nevertheless, T gondii infec-
tion may still cause a very similar clinical
six months old and predominantly have signs of an ascending hindleg picture in dogs and must be included
paralysis, with the associated lesions of polyradiculoneuritis and in the differential diagnosis. In addition,
until recently, another coccidian species,
granulomatous polymyositis. Although any organ may be affected, Hammondia heydorni, was not known to
infections are more common in the central nervous system, muscles, be of clinical significance in dogs but is
now emerging as a cause of diarrhoea.
lungs and skin. Ante-mortem diagnosis is difficult but serology and This review attempts to present the cur-
cytology can aid diagnosis. The diagnosis can be confirmed by rent state of knowledge on neosporosis
and hammondiosis in dogs in a concise
histology, immunohistochemistry, the use of molecular techniques form focusing on clinical signs. It discusses
on biopsy material, or on post-mortem examination. Neospora treatment and preventive measures avail-
able to dog owners and veterinarians.
caninum oocysts are rarely found in faeces and must be
differentiated from oocysts of related coccidians such as
Hammondia heydorni and Toxoplasma gondii. Hammondia heydorni LIFE CYCLE
can cause diarrrhoea in immunosuppressed dogs. Neosporosis
Dogs (Canis familiaris) and coyotes (Canis
should be suspected in young pups with an ascending paralysis of latrans) are a definitive host of N caninum
(McAllister and others 1998, Gondim and
the hindlegs. Treatment with clindamycin and potentiated
others 2004). Experimentally and natu-
sulphonamides may be useful in cases where muscular atrophy rally infected dogs shed low numbers of
oocysts, with oocyst excretion commen-
and fibrosis are absent. Feeding of raw meat is a potential risk
cing between five and 13 days postinfec-
factor for infection of dogs and should be discouraged. tion and shedding lasting for up to 27
days. Oocysts sporulate 24 to 72 hours
after being passed in the faeces and then
become infective. Recent reports suggest
M. P. REICHEL, J. T. ELLIS* AND INTRODUCTION that young puppies are more likely to shed
J. P. DUBEYy more oocysts than older dogs (Gondim
A neosporosis-like illness was first recog- and others 2005). In addition, the source
Journal of Small Animal Practice (2007)
48, 308–312 nised in Norway in boxer dogs which of infective material is important, as
DOI: 10.1111/j.1748-5827.2006.00236.x developed neurological disorders soon oocyst production is far greater in dogs
after birth (Bjerkås and others 1984), but fed N caninum-infected beef than in those
the organism, Neospora caninum, causing fed infected-murine tissues (Gondim and
the disease was not identified until it was others 2002).
named a few years later (Dubey and others There is epidemiological evidence that
1988). Since then, infections due to N can- the life cycle can be maintained between
Gribbles Veterinary Pathology, 840 Tremaine inum have been described in many coun- dogs and cattle. Dogs have been infected
Avenue, Palmerston North, New Zealand tries worldwide (Dubey and Lindsay 1996, by feeding infected bovine foetal mem-
*Department of Medical and Molecular Lindsay and Dubey 2000, Dubey and branes (Dijkstra and others 2001), and
Biosciences, University of Technology, Sydney, others 2002). it is believed that infection may occur
PO Box 123, Broadway, Sydney, New South
Wales 2007, Australia
Before the identification of N caninum from ingested raw meat such as beef that
as a new species, and the realisation that contain the encysted stage of the parasite
yAnimal Parasitic Diseases Laboratory, ANRI,
ARS, U.S. Department of Agriculture, BARC, neosporosis was a separate disease, dogs (Peters and others 2001). Infection of
Building 1001, Beltsville, MD 20705-2350, USA were often diagnosed as suffering from dogs as a primary host produces infective

308 Journal of Small Animal Practice
Vol 48
June 2007
Ó 2007 British Small Animal Veterinary Association
 Neosporosis and hammondiosis in dogs

application of molecular techniques (Šlapeta

and others 2002a). Similar to N caninum,
H heydorni has a dog-cattle life cycle
(Dubey and others 2002), although other
animals (such as deer) can act as interme-
diate hosts (Dubey and others 2004b).

SEROPREVALENCE

The prevalence of infection with N can-

inum in clinically normal dog populations
ranges from apparently nil in the Falkland
Islands and Kenya to around 20 per cent
of a population being infected in Tanzania
and Uruguay (Barber and others 1997a)
and up to 54
2 per cent in dogs from beef
farms in Argentina (Basso and others
2001). Similar levels of seroprevalence are
reported from other parts of South Amer-
ica (de Souza and others 2002, Gennari
and others 2002, Fernandes and others
2004, Azevedo and others 2005). World-
wide prevalences have been summarised
(Lindsay and Dubey 2000, Dubey 2003).
In Europe, two to 13 per cent of ran-
domly studied dogs were seropositive to
N caninum in Austria (Wanha and others
2005), Belgium (Barber and others 1997b),
England (Trees and others 1993), Hungary
(Hornok and others 2006), Italy (Capelli
and others 2004), the Netherlands (Wouda
and others 1999), Spain (Ortuño and
FIG 1. Life cycle of Neospora caninum. The dog is recognised as a definitive host (1), and oocysts others 2002), Sweden (Björkman and others
are shed unsporulated in its faeces. Sporulation of oocysts occurs outside the host. Sporulated 1994) and Turkey (Coskun and others
oocysts can infect cattle orally. N. caninum is thought to persist in the brain and central nervous
system of cattle and a number of other intermediate hosts, such as sheep, goats, deer and water 2000). In selected subpopulations, the sero-
buffalo. During pregnancy, the parasite may be transmitted transplacentally at high efficiency (2) to prevalence may be considerably higher
calves. If this occurs during the first trimester, abortion typically occurs (3). Transplacental or lower (Basso and others 2001, Antony
transmission is the most important method of transmission, evidence, however, shows that
maintenance of infection of a herd requires postnatal infection to occur as well. Transmission to and Williamson 2003, Wanha and others
a definitive host via adult or fetal fluids or tissues must, therefore, occur (4) 2005). For example, higher seroprevalences
have been reported for farm dogs than
for urban dogs (Sawada and others 1998,
oocysts, which have the potential to infect Transplacental transmission from dam
Basso and others 2001, Sager and others
cattle (Fig 1). N caninum oocysts have, in to pup appears to be efficient but is
2006).
turn, been shown to be infective to cattle unlikely to sustain infection in the dog
experimentally (De Marez and others population (Barber and Trees 1998,
1999, Trees and others 2002). Adult cattle Dubey and others 2005). Up to 50 per
generally show no signs of clinical disease cent of pups of N caninum-positive dams COPROSCOPIC STUDIES OF
but may abort sometimes in epidemic pro- might become infected transplacentally, INFECTION
portions. Transplacental infection in cattle, with 25 per cent developing clinical signs.
however, is very efficient and up to 80 to 90 In contrast, little is known about Oocysts of N caninum, which are found
per cent of calves might be born infected, H heydorni, which has frequently been infrequently in the faeces of dogs, are also
yet clinically normal if the dam is infected confused in the past with N caninum, indistinguishable from those of some
with N caninum in the later stages (last tri- although both are now recognised as other apicomplexan parasites that cycle
mester) of gestation (Paré and others 1996, independent species (Dubey and others through dogs, in particular H heydorni
Williams and others 2000). 2002) and can be distinguished by the (Lindsay and others 1999b, Šlapeta and

Journal of Small Animal Practice
Vol 48
June 2007
Ó 2007 British Small Animal Veterinary Association 309

others 2002a,b, Ellis and Pomroy 2003).
Oocysts of the feline coccidians, T gondii
and Hammondia hammondi, can also be
mechanically transmitted by dogs and
hence either may be present in the faeces
of dogs (Lindsay and others 1997, Schares
and others 2005); apparently some of
these oocysts pass through the canine
gut unexcysted after dogs ingest cat faeces.
Hence, oocyst morphology is not helpful
in species identification of these cyst-form-
ing coccidian found in faeces. Molecular
techniques, based on polymerase chain
reaction (PCR) and sequencing, can help
identify the species present; however, these
approaches are constrained by the
number of oocysts present. Infection of
gerbils with oocysts can result in serocon-
version if N caninum is present (Schares
and others 2005); however, it is difficult
to determine the presence of H heydorni
because of the absence of appropriate sero-
logical tests.
Studies have reported diarrhoea in
normal and an immunosuppressed dog
associated with H heydorni infection
(Blagburn and others 1988, Schares and
others 2005, Abel and others 2006). Dogs
can repeatedly shed oocysts of N caninum
(McGarry and others 2003, Gondim and
others 2005, Sager and others 2006).
There are a number of possible explana-
tions for these observed phenomena, such
as failure to induce host immunity; how-
ever, the results of research on this topic
have yet to be reported.
SOURCES OF INFECTION
Dogs consuming raw meat are at risk of
becoming infected with both N caninum
(McAllister and others 1998, Lindsay
and others 1999a) and H heydorni (Dubey
and others 2003, Schares and others
2005). A wide range of animals, known
to act as intermediate hosts for these para-
sites, may be infected and these include
cattle, sheep, and deer, among others
(Dubey 2003, Rosypal and Lindsay
2005). Evidence for the importance of
raw meat in the transmission of N caninum
to dogs is provided by the higher
seroprevalance among hunting hounds
(51 per cent) compared with pet dogs
(7
5 per cent) living in the UK (Trees
310
M. P. Reichel and others
and Williams in [Hemphill and Gottstein
2000]).
CLINICAL SIGNS
Two main neurological forms are ob-
served: encephalomyelitis and myositis-
polyradiculoneuritis. Protozoal myositis-
polyradiculoneuritis is probably the most
commonly reported infectious myositis
in dogs. It is generally observed in dogs
under six months of age, which were
infected transplacentally, and over half
of these dogs may go on to have signs of
a progressive, ascending paralysis of the
hindlegs (Fig 2). Muscle atrophy and
a rigid hyperextension of the limb often
develops (Barber and Trees 1996). Serum
creatine kinase levels may be increased.
Muscle biopsy is indicated and may
reveal non-suppurative inflammation and
tachyzoites within myocytes (Knowler
and Wheeler 1995, Ruehlmann and others
1995).
Neurological signs associated with pro-
tozoal encephalomyelitis are variable and
may reflect a focal or multi-focal disease
process. Cerebrospinal fluid exhibits mild
mixed pleocytosis and protein increase.
Eosinophils are found in only a few cases.
Adults are more commonly affected than
the young. N caninum has a predilection
for lumbosacral roots in puppies, result-
FIG 2. Boxer pup with fixated hindleg due to
neosporosis (reproduced with permission from
the New Zealand Veterinary Journal)
Journal of Small Animal Practice
Vol 48
June 2007
Ó 2007 British Small Animal Veterinary Association
ing in pelvic limb atrophy and immobile
joints (contractures) (Ruehlmann and
others 1995).
There are also reports of dermatitis,
myocarditis and pneumonia (Ruehlmann
and others 1995, McInnes and others
2006). No reports of sex predilection exist
but the boxer breed is frequently men-
tioned (Trees and others 1993, Barber
and others 1997b, Cringoli and others
2002). Most affected dogs come from lit-
ters where other littermates are infected,
but which show no signs of clinical infec-
tion (Dubey and others 1988, 2004a
Dubey and Lindsay 1996, Reichel and
others 1998).
A parasitic dermatitis containing N
caninum-like tachyzoites is increasingly being
recognised (McInnes and others 2006).
Other organs infected might include heart,
lung and, less frequently, liver, rarely the
adrenal gland, thyroid gland and uterus,
although clinical signs resulting from dam-
age to these organs are rarely reported.
Hammondia heydorni is not normally
associated with disease in dogs, although
it is increasingly becoming associated with
diarrhoea (Schares and others 2005, Abel
and others 2006).
DIAGNOSIS
Dogs typically seroconvert two to three
weeks after infection with N caninum
(McAllister and others 1998), and so diag-
nosis of neosporosis in the live animal
can be based on clinical signs with sero-
logy. Antibody titres in the indirect fluore-
scent antibody test rarely exceed 1:800 in
clinically unaffected dogs (Barber and
Trees 1996). However, dogs with proven
neosporosis may have only low antibody
titres (Dubey and others 1998, 2005).
Immunoglobulin M detection may not
be rewarding, especially in congenitally
infected dogs (Dubey and others 1998).
A commercially available competitive
ELISA test is also available (and validated)
for testing dog sera (Capelli and others
2006).
Histological examinations (from
biopsy material or post-mortem samples)
yield typical lesions of meningoencephalo-
myelitis and myositis in various muscles
throughout the body (Barber and others

1996). Molecular tools like PCR can
also help in providing a diagnosis (Ellis
1998).
Identification of the disease-causing
organism can be made by its isolation into
cell cultures or immunosuppressed mice,
immunohistochemistry or molecular tech-
niques (Dubey and others 1998, 2004a).
Numerous N caninum tachyzoites may
be found in fluid squeezed from dermal
lesions (Dubey and others 1988). For
unknown reasons, there is immunohisto-
chemical cross-reactivity between N can-
inum and T gondii in dermal neosporosis.
TREATMENT
Early treatment with potentiated sulpho-
namides and clindamycin may resolve
clinical disease, as described in 10 of 16
cases in one report from the UK (Barber
and Trees 1996). Clindamycin was used
at dose rates of up to 22 mg/kg twice a
day for up to 60 days, 15 to 30 mg/kg
potentiated sulphonamides twice a day
for up to 28 days, alone or in combi-
nation with 1 mg/kg pyrimethamine
daily for up to 35 days. Treatment with
clindamycin is very effective in treating
cutaneous neosporosis (Dubey and others
1995). Frequently, however, the limb
lesions progress to muscle atrophy, and
in about 50 per cent of cases rigid hyper-
extension develops that does not respond
to treatment (Fig 2). Ponazuril, at 20 mg/kg,
has been shown to be highly efficacious in
treating experimentally induced neospo-
rosis in mice and cattle, and might be use-
ful in dogs as well (Gottstein and others
2001, Kritzner and others 2002). Treat-
ment schedule and doses are given by
Dubey and Lappin (2006).
PREVENTION
There are no drugs or vaccines that
will prevent neosporosis in dogs. Feeding
of uncooked beef and other animal
tissues that may be infected with N cani-
num to dogs is an important route of
transmission. Feeding of raw meat to
dogs, especially where they are close to
susceptible cattle should, therefore, be
stopped.
Journal of Small Animal Practice
Vol 48
June 2007
Ó 2007 British Small Animal Veterinary Association
Neosporosis and hammondiosis in dogs
CLOSING REMARKS
Neospora caninum infection is prevalent
in dog populations around the world;
clinical disease in the form of an ascend-
ing hindleg paralysis, however, is much
less frequently reported. In comparison,
H heydorni is an emerging pathogen in
dogs and should be considered in cases
of diarrhoea.
Dogs should also be prohibited from
access to aborted bovine foetal material,
including foetal membranes, birth fluids
and dead calves. One suggested means
of achieving this is to control exposure
of working dogs on farms by muzzling
(Antony and Williamson 2001).
Transplacental transmission occurs in
dogs, as it does in cattle and is reported
to occur repeatedly in series of litters from
an infected dam. It may be advisable to
cease breeding from infected dams, or
to introduce a serological monitoring pro-
gramme for the young pups of such
dams, to facilitate early (even prophylactic)
intervention before the onset of clinical
signs.
Treatment with potentiated sulphona-
mides and clindamycin promises success if
begun before atrophy and fibrotic lesions
have occurred.
Acknowledgements
Our thanks to Cindy Burton, whose story
prompted us to complete this article
(http://www.angelfire.com/stars3/starridge/
Gatsby_Mtarni.htm). Thanks are also due
to Dr Laurent Garosi for valuable com-
ments on the manuscript.
References
ABEL, J., SCHARES, G., ORZESZKO, K., GASSER, R. B. &
ELLIS, J. T. (2006) Hammondia isolated from dogs
and foxes are genetically distinct. Parasitology
132, 187-192
ANTONY, A. & WILLIAMSON, N. B. (2001) Recent
advances in understanding the epidemiology of
Neospora caninum in cattle. New Zealand
Veterinary Journal 49, 42-47
ANTONY, A. & WILLIAMSON, N. B. (2003) Prevalence of
antibodies to Neospora caninum in dogs of rural or
urban origin in central New Zealand. New Zealand
Veterinary Journal 51, 232-237
AZEVEDO, S. S., BATISTA, C. S., VASCONCELLOS, S. A.,
AGUIAR, D. M., RAGOZO, A. M., RODRIGUES, A. A.,
ALVES, C. J. & GENNARI, S. M. (2005) Seroepidemi-
ology of Toxoplasma gondii and Neospora
caninum in dogs from the state of Paraiba, North-
east region of Brazil. Research in Veterinary
Science 79, 51-56
BARBER, J. S., GASSER, R. B., ELLIS, J., REICHEL, M. P.,
MCMILLAN, D. & TREES, A. J. (1997a) Prevalence
of antibodies to Neospora caninum in different
canid populations. Journal of Parasitology 83,
1056-1058
BARBER, J. S., PAYNE-JOHNSON, C. E. & TREES, A. J. (1996)
Distribution of Neospora caninum within the cen-
tral nervous system and other tissues of six dogs
with clinical neosporosis. Journal of Small Animal
Practice 37, 568-574
BARBER, J. S. & TREES, A. J. (1996) Clinical aspects of
27 cases of neosporosis in dogs. Veterinary
Record 139, 439-443
BARBER, J. S. & TREES, A. J. (1998) Naturally occurring
vertical transmission of Neospora caninum in
dogs. International Journal for Parasitology 28,
57-64
BARBER, J. S., VAN HAM, L., POLIS, I. & TREES, A. J.
(1997b) Seroprevalence of antibodies to Neo-
spora caninum in Belgian dogs. Journal of Small
Animal Practice 38, 15-16
BASSO, W., VENTURINI, L., VENTURINI, M. C., MOORE, P.,
RAMBEAU, M., UNZAGA, J. M., CAMPERO, C., BACIGALUPE,
D. & DUBEY, J. R. (2001) Prevalence of Neospora
caninum infection in dogs from beef-cattle farms,
dairy farms, and from urban areas of Argentina.
Journal of Parasitology 87, 906-907
BJERKÅS, I., MOHN, S. F. & PRESTHUS, J. (1984) Uniden-
tified cyst-forming sporozoon causing encephalo-
myelitis and myositis in dogs. Zeitschrift für
Parasitenkunde 70, 271-274
BJÖRKMAN, C., LUNDEN, A. & UGGLA, A. (1994) Preva-
lence of antibodies to Neospora caninum and
Toxoplasma gondii in Swedish dogs. Acta Veter-
inaria Scandinavica 35, 445-447
BLAGBURN, B. L., LINDSAY, D. S., SWANGO, L. J., PIDGEON,
G. L. & BRAUND, K. G. (1988) Further character-
ization of the biology of Hammondia heydorni .
Veterinary Parasitology 27, 193-198
CAPELLI, G., NARDELLI, S., DI REGALBONO, A. F., SCALA, A. &
PIETROBELLI, M. (2004) Sero-epidemiological
survey of Neospora caninum infection in dogs
in north-eastern Italy. Veterinary Parasitology
123, 143-148
CAPELLI, G., NATALE, A., NARDELLI, S., FRANGIPANE DI
REGALBONO, A. & PIETROBELLI, M. (2006) Validation
of a commercially available cELISA test for canine
neosporosis against an indirect fluorescent anti-
body test (IFAT). Preventive Veterinary Medicine
73, 315-320
COSKUN, S. Z., AYDYN, L. & BAUER, C. (2000) Sero-
prevalence of Neospora caninum infection in
domestic dogs in Turkey. Veterinary Record
146, 649
CRINGOLI, G., RINALDI, L., CAPUANO, F., BALDI, L.,
VENEZIANO, V. & CAPELLI, G. (2002) Serological
survey of Neospora caninum and Leishmania
infantum co-infection in dogs. Veterinary Parasi-
tology 106, 307-313
DE MAREZ, T., LIDDELL, S., DUBEY, J. P., JENKINS, M. C. &
GASBARRE, L. (1999) Oral infection of calves with
Neospora caninum oocysts from dogs: humoral
and cellular immune responses. International
Journal for Parasitology 29, 1647-1657
DE SOUZA, S. L., GUIMARAES, J. S. Jr., FERREIRA, F., DUBEY,
J. P. & GENNARI, S. M. (2002) Prevalence of Neo-
spora caninum antibodies in dogs from dairy
cattle farms in Parana, Brazil. Journal of
Parasitology 88, 408-409
DIJKSTRA, T., EYSKER, M., SCHARES, G., CONRATHS, F. J.,
WOUDA, W. & BARKEMA, H. W. (2001) Dogs shed
Neospora caninum oocysts after ingestion of
naturally infected bovine placenta but not after
ingestion of colostrum spiked with Neospora
caninum tachyzoites. International Journal for
Parasitology 31, 747-752
DUBEY, J. P. (2003) Review of Neospora caninum and
neosporosis in animals. Korean Journal of
Parasitology 41, 1-16
DUBEY, J. P., BARR, B. C., BARTA, J. R., BJERKAS, I.,
BJORKMAN, C., BLAGBURN, B. L., BOWMAN, D. D., BUXTON,
D., ELLIS, J. T., GOTTSTEIN, B., HEMPHILL, A., HILL, D. E.,
HOWE, D. K., JENKINS, M. C., KOBAYASHI, Y., KOUDELA,
B., MARSH, A. E., MATTSSON, J. G., MCALLISTER, M. M.,
MODRY, D., OMATA, Y., SIBLEY, L. D., SPEER, C. A.,
311

TREES, A. J., UGGLA, A., UPTON, S. J., WILLIAMS, D. J. &
LINDSAY, D. S. (2002) Redescription of Neospora
caninum and its differentiation from related coc-
cidia. International Journal for Parasitology 32,
929-946
DUBEY, J. P., CARPENTER, J. L., SPEER, C. A., TOPPER, M. J. &
UGGLA, A. (1988) Newly recognized fatal protozoan
disease of dogs. Journal of the American Veterinary
Medical Association 192, 1269-1285
DUBEY, J. P., DOROUGH, K. R., JENKINS, M. C., LIDDELL, S.,
SPEER, C. A., KWOK, O. C. & SHEN, S. K. (1998)
Canine neosporosis: clinical signs, diagnosis,
treatment and isolation of Neospora caninum in
mice and cell culture. International Journal for
Parasitology 28, 1293-1304
DUBEY, J. P., KNICKMAN, E. & GREENE, C. E. (2005) Neo-
natal Neospora caninum infections in dogs. Acta
Parasitologica 50, 176-179
DUBEY, J. P. & LAPPIN, M. R. (2006) Toxoplasmosis
and neosporosis. In: Infectious Diseases of the
Dog and Cat. 3rd edn. Ed C. E. Greene. Saunders
Elsevier, St Louis, MO. pp 754-775
DUBEY, J. P. & LINDSAY, D. S. (1996) A review of
Neospora caninum and neosporosis. Veterinary
Parasitology 67, 1-59
DUBEY, J. P., METZGER, F. L. Jr., HATTEL, A. L., LINDSAY,
D. S. & FRITZ, D. L. (1995) Canine cutaneous neo-
sporosis: clinical improvement with clindamycin.
Veterinary Dermatology 6, 37-43
DUBEY, J. P., ROSS, A. D. & FRITZ, D. (2003) Clinical
Toxoplasma gondii, Hammondia heydorni, and
Sarcocystis spp. infections in dogs. Parassito-
logia 45, 141-146
DUBEY, J. P., SREEKUMAR, C., KNICKMAN, E., MISKA, K. B.,
VIANNA, M. C., KWOK, O. C., HILL, D. E., JENKINS, M. C.,
LINDSAY, D. S. & GREENE, C. E. (2004a) Biologic,
morphologic, and molecular characterisation
of Neospora caninum isolates from littermate
dogs. International Journal for Parasitology 34,
1157-1167
DUBEY, J. P., SREEKUMAR, C., MISKA, K. B., HILL, D. E.,
VIANNA, M. C. & LINDSAY, D. S. (2004b) Molecular
and biological characterization of Hammondia
heydorni-like oocysts from a dog fed hearts from
naturally infected white-tailed deer (Odocoileus
Virginianus). The Journal of Parasitology 90,
1174-1176
ELLIS, J. T. (1998) Polymerase chain reaction ap-
proaches for the detection of Neospora caninum
and Toxoplasma gondii . International Journal for
Parasitology 28, 1053-1060
ELLIS, J. T. & POMROY, W. E. (2003) Hammondia hey-
dorni in the faeces of a greyhound. New Zealand
Veterinary Journal 51, 38-39
FERNANDES, B. C., GENNARI, S. M., SOUZA, S. L., CARVALHO,
J. M., OLIVEIRA, W. G. & CURY, M. C. (2004) Pre-
valence of anti-Neospora caninum antibodies in
dogs from urban, periurban and rural areas of
the city of Uberlandia, Minas Gerais–Brazil. Veter-
inary Parasitology 123, 33-40
GENNARI, S. M., YAI, L. E., D’AURIA, S. N., CARDOSO,
S. M., KWOK, O. C., JENKINS, M. C. & DUBEY, J. P.
(2002) Occurrence of Neospora caninum anti-
bodies in sera from dogs of the city of Sao Paulo,
Brazil. Veterinary Parasitology 106, 177-179
GONDIM, L. F., GAO, L. & MCALLISTER, M. M. (2002)
Improved production of Neospora caninum
oocysts, cyclical oral transmission between dogs
and cattle, and in vitro isolation from oocysts.
Journal of Parasitology 88, 1159-1163
312
M. P. Reichel and others
GONDIM, L. F., MCALLISTER, M. M. & GAO, L. (2005)
Effects of host maturity and prior exposure history
on the production of Neospora caninum oocysts
by dogs. Veterinary Parasitology 134, 33-39
GONDIM, L. F., MCALLISTER, M. M., PITT, W. C. &
ZEMLICKA, D. E. (2004) Coyotes (Canis latrans)
are definitive hosts of Neospora caninum . Inter-
national Journal for Parasitology 34, 159-161
GOTTSTEIN, B., EPERON, S., DAI, W. J., CANNAS, A., HEMPHILL,
A. & GREIF, G. (2001) Efficacy of toltrazuril and
ponazuril against experimental Neospora caninum
infection in mice. Parasitology Research 87, 43-48
HEMPHILL, A. & GOTTSTEIN, B. (2000) A European per-
spective on Neospora caninum . International
Journal for Parasitology 30, 877-924
HORNOK, S., EDELHOFER, R., FOK, E., BERTA, K., FEJES, P.,
REPASI, A. & FARKAS, R. (2006) Canine neosporosis
in Hungary: screening for seroconversion of house-
hold, herding and stray dogs. Veterinary Parasitol-
ogy 137, 197-201
KNOWLER, C. & WHEELER, S. J. (1995) Neospora can-
inum infection in three dogs. Journal of Small Ani-
mal Practise 36, 172-177
KRITZNER, S., SAGER, H., BLUM, J., KREBBER, R., GREIF, G. &
GOTTSTEIN, B. (2002) An explorative study to assess
the efficacy of toltrazuril-sulfone (Ponazuril) in
calves experimentally infected with Neospora
caninum . Annals of Clinical Microbiology and
Antimicrobials 1, 4
LINDSAY, D. S. & DUBEY, J. P. (2000) Canine neosporo-
sis . Journal of Veterinary Parasitology 14, 1-11
LINDSAY, D. S., DUBEY, J. P., BUTLER, J. M. & BLAGBURN,
B. L. (1997) Mechanical transmission of Toxo-
plasma gondii oocysts by dogs. Veterinary Parasi-
tology 73, 27-33
LINDSAY, D. S., DUBEY, J. P. & DUNCAN, R. B. (1999a)
Confirmation that the dog is a definitive host for
Neospora caninum . Veterinary Parasitology 82,
327-333
LINDSAY, D. S., UPTON, S. J. & DUBEY, J. P. (1999b) A
structural study of the Neospora caninum oocyst.
International Journal for Parasitology 29, 1521-
1523
MCALLISTER, M. M., DUBEY, J. P., LINDSAY, D. S., JOLLEY,
W. R., WILLS, R. A. & MCGUIRE, A. M. (1998) Dogs
are definitive hosts of Neospora caninum . Inter-
national Journal for Parasitology 28, 1473-1478
MCGARRY, J. W., STOCKTON, C. M., WILLIAMS, D. J. &
TREES, A. J. (2003) Protracted shedding of oocysts
of Neospora caninum by a naturally infected fox-
hound. The Journal of Parasitology 89, 628-630
MCINNES, L. M., IRWIN, P., PALMER, D. G. & RYAN, U. M.
(2006) In vitro isolation and characterisation of
the first canine Neospora caninum isolate in Aus-
tralia. Veterinary Parasitology 137, 355-363
MUNDAY, B. L., DUBEY, J. P. & MASON, R. W. (1990)
Neospora caninum infection in dogs. Australian
Veterinary Journal 67, 76-77
ORTUÑO, A., CASTELLA, J. & ALMERIA, S. (2002) Seropre-
valence of antibodies to Neospora caninum in
dogs from Spain. Journal of Parasitology 88,
1263-1266
PARÉ, J., THURMOND, M. C. & HIETALA, S. K. (1996) Con-
genital Neospora caninum infection in dairy cattle
and associated calfhood mortality. Canadian Jour-
nal of Veterinary Research 60, 133-139
PATITUCCI, A. N., ALLEY, M. R., JONES, B. R. & CHARLESTON,
W. A. G. (1997) Protozoal encephalomyelitis
of dogs involving Neospora caninum and Toxo-
plasma gondii in New Zealand. New Zealand
Veterinary Journal 45, 231-235
Journal of Small Animal Practice
Vol 48
June 2007
Ó 2007 British Small Animal Veterinary Association
PETERS, M., LUTKEFELS, E., HECKEROTH, A. R. & SCHARES, G.
(2001) Immunohistochemical and ultrastructural
evidence for Neospora caninum tissue cysts in
skeletal muscles of naturally infected dogs and
cattle. International Journal for Parasitology 31,
1144-1148
REICHEL, M. P., THORNTON, R. N., MORGAN, P. L., MILLS,
R. J. M. & SCHARES, G. (1998) Neosporosis in a
pup. New Zealand Veterinary Journal 46, 106-110
ROSYPAL, A. C. & LINDSAY, D. S. (2005) The sylvatic
cycle of Neospora caninum: where do we go from
here? Trends in Parasitology 21, 439-440
RUEHLMANN, D., PODELL, M., OGLESBEE, M. & DUBEY, J. P.
(1995) Canine neosporosis: a case report and lit-
erature review. Journal of the American Animal
Hospital Association 31, 174-183
SAGER, H., MORET, C. S., MULLER, N., STAUBLI, D., ESPOSITO,
M., SCHARES, G., HASSIG, M., STARK, K. & GOTTSTEIN, B.
(2006) Incidence of Neospora caninum and
other intestinal protozoan parasites in populations
of Swiss dogs. Veterinary Parasitology 139,
84-92
SAWADA, M., PARK, C. H., KONDO, H., MORITA, T., SHIMADA,
A., YAMANE, I. & UMEMURA, T. (1998) Serological
survey of antibody to Neospora caninum in Japa-
nese dogs. Journal of Veterinary Medical Science
60, 853-854
SCHARES, G., PANTCHEV, N., BARUTZKI, D., HEYDORN, A. O.,
BAUER, C. & CONRATHS, F. J. (2005) Oocysts of
Neospora caninum, Hammondia heydorni, Toxo-
plasma gondii and Hammondia hammondi in
faeces collected from dogs in Germany. Interna-
tional Journal for Parasitology 35, 1525-1537
ŠLAPETA, J. R., KOUDELA, B., VOTYPKA, J., MODRÝ, D.,
HOREJŠ, R. & LUKEŠ, J. (2002a) Coprodiagnosis
of Hammondia heydorni in dogs by PCR based
amplification of ITS 1 rRNA: differentiation from
morphologically indistinguishable oocysts of
Neospora caninum . Veterinary Journal (London,
England: 1997) 163, 147-154
ŠLAPETA, J. R., MODRÝ, D., KYSELOVÁ, I., HOREJŠ, R., LUKEŠ,
J. & KOUDELA, B. (2002b) Dog shedding oocysts of
Neospora caninum: PCR diagnosis and molecular
phylogenetic approach. Veterinary Parasitology
109, 157-167
TREES, A. J., GUY, F., TENNANT, B. J., BALFOUR, A. H. &
DUBEY, J. P. (1993) Prevalence of antibodies to
Neospora caninum in a population of urban dogs
in England. Veterinary Record 132, 125-126
TREES, A. J., MCALLISTER, M. M., GUY, C. S., MCGARRY,
J. W., SMITH, R. F. & WILLIAMS, D. J. (2002) Neospora
caninum: oocyst challenge of pregnant cows. Vet-
erinary Parasitology 109, 147-154
WANHA, K., EDELHOFER, R., GABLER-EDUARDO, C. & PROSL, H.
(2005) Prevalence of antibodies against Neo-
spora caninum and Toxoplasma gondii in dogs
and foxes in Austria. Veterinary Parasitology 128,
189-193
WILLIAMS, D. J., GUY, C. S., MCGARRY, J. W., GUY, F.,
TASKER, L., SMITH, R. F., MACEACHERN, K., CRIPPS,
P. J., KELLY, D. F. & TREES, A. J. (2000) Neospora
caninum-associated abortion in cattle: the time
of experimentally-induced parasitaemia during
gestation determines foetal survival. Parasitol-
ogy 121, 347-358
WOUDA, W., DIJKSTRA, T., KRAMER, A. M., VAN MAANEN, C. &
BRINKHOF, J. M. (1999) Seroepidemiological evi-
dence for a relationship between Neospora cani-
num infections in dogs and cattle. International
Journal for Parasitology 29, 1677-1682