Besnoitiosis is a chronic parasitic disease that mainly affects ungulates. The disease is characterized initially by fever and then edema, which progresses to hyperkeratosis, scleroderma with skin folding, and tissue cysts in subcutaneous tissue and mucous membranes. Serological testing is key for diagnosis and control. No treatment options or vaccines are available.
Besnoitiosis is caused by tissue cyst–forming apicomplexan parasites of the genus Besnoitia. The disease, which mainly affects ungulates, is usually chronic and debilitating, and it is characterized by skin lesions and systemic clinical signs of varying severity. Besnoitiosis compromises animal welfare and is responsible for severe economic losses: infected cattle may lose weight, dams may abort during the febrile stage and have decreased milk production, males may become infertile or even sterile, and the hides of affected animals are of decreased value for leather production.
The skin lesions and visible scleral tissue cysts characteristic of besnoitiosis are similar among affected ungulates. There are no effective treatments. For control in cattle, the only permitted vaccine worldwide is a live vaccine used to immunize imported bulls in Israel. Thus, control relies entirely on accurate diagnosis and herd management measures.
Etiology of Besnoitiosis
The genus Besnoitia belongs to a group of cyst-forming coccidians (subphylum Apicomplexa, family Sarcocystidae) related to Toxoplasma gondii and Neospora caninum. Besnoitia comprises 10 recognized species: B akodoni, B bennetti, B besnoiti, B caprae, B darlingi, B jellisoni, B neotomofelis, B oryctofelisi, B tarandi, and B wallacei.
Several of these species have both definitive and intermediate hosts (heteroxenous life cycle). However, the identity of the definitive host remains unknown for B besnoiti, B tarandi, B bennetti, and B caprae, which infect ungulates (bovids, wild ruminants, equids, and goats, respectively) that act as intermediate hosts. The definitive hosts are also unknown for B akodoni and B jellisoni, which infect rodents.
The cheetah (Acinonyx jubatus) has been proposed as the definitive host of an undescribed Besnoitia species (a B darlingi–like species termed B acinonyx) (1) whose natural intermediate hosts are unknown.
The table summarizes the species included in the Besnoitia genus and their intermediate hosts (ungulates, rodents, lagomorphs, marsupials, or reptiles), definitive hosts (if known), and geographical distributions.
Natural Hosts and Geographical Distribution of Besnoitia Species
Species | Intermediate Hosts | Definitive Hosts | Geographical Distribution |
|---|---|---|---|
Besnoitia besnoiti | Bovine, antelope (Aepyceros melampus), impala, blue wildebeest (Connochaetes taurinus), red deer (Cervus elaphus), roe deer (Capreolus capreolus) | Unknowna | Africa, Asia, Europe; one case report in Venezuelab |
B bennetti | Donkey (Equus africanus asinus), horse (E ferus caballus), zebra (E zebra, E quagga, E grevyi) | Unknowna | Africa, Asia, US, Europe |
B caprae | Goat (Capra aegagrus hircus) | Unknowna | Iran, Kenya; one isolated case report in Nigeriac |
B tarandi | Musk ox (Ovibos moschatus), reindeer (Rangifer tarandus), mule deer (Odocoileus hemionus) | Unknowna | Arctic region (Canada, Finland, Russia, Sweden) |
B akodoni | Akodon montensis (rodent) | Unknown | Brazild |
B jellisoni | Dipodomys spp (rodent), Peromyscus maniculatus (rodent) | Unknown | US |
B darlingi | Ameiva spp (reptile), Basiliscus spp (reptile), Didelphis marsupialis (marsupial) | Cat (Felis catus) | US, Panama |
B neotomofelis | Neotoma micropus (rodent) | Cat (Felis catus) | USe |
B oryctofelisi | Oryctolagus cuniculus (lagomorph) | Cat (Felis catus) | Argentinaf |
B wallacei | Rattus exulans (rodent) | Cat (Felis catus) | Australia, Hawaii, Japan, Kenya |
a A carnivorous definitive host is suspected for B besnoiti. One study suggested the role of felids as definitive hosts (Peteshev VM, Galuzo IG, Polomoshnov AP. Cats—definitive hosts of Besnoitia [Besnoitia besnoiti]. Izvest Akad Nauk Kazakh Ser Biol. 1974;1:33-38). However, extensive studies carried out by other researchers have been inconclusive (Diesing L, Heydorn AO, Matuschka FR, et al. Besnoitia besnoiti: studies on the definitive host and experimental infections in cattle. Parasitol Res. 1988;75(2):114-117. doi:10.1007/BF00932710. Basso W, Schares G, Gollnick NS, Rütten M, Deplazes P. Exploring the life cycle of Besnoitia besnoiti—experimental infection of putative definitive and intermediate host species. Vet Parasitol. 2011;178(3-4):223-234. doi:10.1016/j.vetpar.2011.01.027). b Vogelsang EG, Gallo P. Globidium besnoiti (Marotel, 1912) y habronemosis cutánea en bovinos de Venezuela. Rev Med Vet Parasit Caracas. 1941;3:153-155. c Sambo SJ, Ibrahim ND, Esievo KA, Hambolu JO, Makoshi MS, Tekdek LB. First incidence of caprine besnoitiosis in Kaduna State of Nigeria. J Anim Vet Adv. 2007;6:673-675. d Dubey JP, Sreekumar C, Rosenthal BM, Lindsay DS, Grisard EC, Vitor RW. Biological and molecular characterization of Besnoitia akodoni n.sp. (Protozoa: Apicomplexa) from the rodent Akodon montensis in Brazil. Parassitologia. 2003;45(2):61-70. e Dubey JP, Yabsley MJ. Besnoitia neotomofelis n. sp. (Protozoa: Apicomplexa) from the southern plains woodrat (Neotoma micropus). Parasitology. 2010;137(12):1731-1747. doi:10.1017/S0031182010000636 f Dubey JP, Sreekumar C, Lindsay DS, et al. Besnoitia oryctofelisi n. sp. (Protozoa: Apicomplexa) from domestic rabbits. Parasitology. 2003;126(Pt 6):521-539. | |||
Intermediate hosts harbor two asexual parasitic stages: tachyzoites and cyst-forming bradyzoites. The former are responsible for the acute stage of besnoitiosis; the latter, for the chronic stage. Upon infection, tachyzoites proliferate mainly in endothelial cells. Next, tachyzoites transform into bradyzoites to evade the host's immune response, and they pack inside thick-walled cysts (see ). These cysts are found mostly in subcutaneous tissues and mucous membranes during the chronic stage of besnoitiosis and are considered pathognomonic of the disease.
Photomicrograph of a cross-section of a tissue cyst surrounded by an inflammatory infiltrate sampled from a blood vessel in the pampiniform plexus of a bull with besnoitiosis. H&E stain; scale bar = 50 mcm.
Courtesy of Dr. Gema Álvarez García.
Epidemiology of Besnoitiosis
The range of bovine besnoitiosis has expanded since the late 1990s in Europe, and the disease is now endemic in France, Spain, Portugal, and northern Italy. Cases of B besnoiti infection in Germany, Switzerland, Italy, Croatia, Hungary, Belgium, and Ireland have also been described (2). Besnoitiosis is widely distributed in sub-Saharan African countries (Angola, Botswana, Cameroon, Egypt, Kenya, Namibia, Nigeria, Tanzania, Uganda, South Africa, Sudan, Swaziland, Zambia, Zimbabwe). In Asia, the disease is present in Israel, and serological or molecular evidence of the infection has been reported in Jordan (3), Turkey (4, 5), Egypt (6, 7, 8), Arabia (9), Iraq (10), and Korea (11). In contrast, case reports have not been documented from North America or Australia.
Besnoitiosis in reindeer, mule deer, and caribou populations is present in Arctic regions of North America and Europe. Moreover, serological evidence of infection has been found in musk oxen and bison (12). Besnoitiosis in goats is restricted to a few countries, such as Iran and Kenya, and one isolated case in Nigeria (13). Finally, besnoitiosis in equids has been reported in France, Belgium, UK, Ireland, Spain, and Italy (14, 15, 16), and the disease is more prevalent in donkeys than previously known and might be emerging in the US (17). Besnotia spp infections in small mammals have mainly been reported in the American continent (18).
It has been postulated that transmission in cattle occurs horizontally, mainly through direct contact between infected and noninfected animals (19) (see ). Tissue cysts can be present in penis mucosa and vestibulum vaginae; these cysts can rupture, and the released bradyzoites can cross mucosal barriers. Accordingly, parasite transmission might be favored by natural breeding. Indeed, breeding a seronegative cow with a seropositive bull increases the infection risk for the cow (20). It has also been reported that a high B besnoitia prevalence leads to infection of bulls within a short time (20).
Cattle and wild ruminants serve as intermediate hosts of Besnoitia besnoiti by harboring two asexual parasite stages: the fast-replicating tachyzoites (responsible for the acute phase of besnoitiosis) and the slow-dividing bradyzoites gathered inside tissue cysts (which are more abundant in the skin and mucous membranes).
Although the identity and relevance of a definitive host for B besnoiti remain unknown, a carnivorous species is suspected; felids are often suggested on the basis of the life cycles of related Besnoitia species: 1) the definitive host ingests tissues infected with tissue cysts; 2) the bradyzoites go through the enteroepithelial cycle that ends with the shedding of oocysts; 3) the oocysts sporulate in the environment and can potentially be ingested by the intermediate host. (Peteshev VM, Galuzo IG, Polomoshnov AP. Cats—definitive hosts of Besnoitia [Besnoitia besnoiti]. Izvest Akad Nauk Kazakh Ser Biol 1974;1:33-38.)
Epidemiological data also support transmission of the parasite in cattle through direct contact (eg, natural breeding), as a consequence of ruptured tissue cysts in genital mucosa and bradyzoites' ability to cross mucous membranes. Only experimental evidence supports mechanical transmission via blood-sucking insects, such as tabanids and stable flies, during periods of high vector activity.
Courtesy of Dr. Gema Álvarez García.
Mechanical transmission of B besnoiti has also been achieved experimentally (21), either with a syringe or with bloodsucking insects (tabanid species and Stomoxyscalcitrans); secretophagous flies have been suggested as potential vectors.
The role of arthropods in parasite transmission is supported by the seasonal incidence of besnoitiosis. Arthropods have a restricted range of transmission, and the parasites they harbor might be either tachyzoites or bradyzoites.
In Europe, roe deer and red deer might act as intermediate hosts of B besnoiti. However, the prevalence of infection in wild ruminants is very low, and these species probably do not play a substantial role in the epidemiology of bovine besnoitiosis. In Africa, in addition to cattle, antelopes and wildebeest are also intermediate hosts.
The definitive host of the Besnoitia spp that infect ungulates is unknown; however, it is suspected to be a carnivore, as with the Besnoitia spp that parasitize small mammals (B wallacei, B darlingi, B neotomofelis, and B oryctofelisi). For the latter species, cats get infected after consuming prey (eg, rodents) harboring tissue cysts; after intestinal replication (similar to that for other coccidia), nonsporulated oocysts are shed in cats' feces. Oocysts sporulate in the environment and are the infectious form for intermediate hosts by the oral route. Neither venereal transmission nor transplacental transmission have been demonstrated for these species in cattle.
Parasite transmission in cattle is favored by risk factors associated with extensive husbandry systems (eg, natural breeding and seasonal exposure to arthropods). Moreover, the movement of cattle with unknown health status and without prior diagnosis of infection upon entry into a new herd has been identified as a key risk factor for besnoitiosis dissemination. Both sexes are equally affected. An increase of serological and clinical prevalence with age has been reported, and clinical prevalence is higher in animals 2–4 years old (22). Clinical cases are sporadic in animals < 1 year old (23).
Pearls & Pitfalls
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In endemic areas, the number of animals showing clinical signs of besnoitiosis is typically low (1–10%) (22); however, the proportion of subclinically infected cattle (parasite carriers) might be high. In these cases, clinical signs are mild and associated with a low parasite load. Severe cases are generally restricted to naive animals recently introduced into an infected herd. In contrast, during epidemic outbreaks, both the incidence rate and the number of clinically affected animals tend to be substantially higher.
Clinical Findings of Besnoitiosis
Once cattle become infected with B besnoiti, the incubation period can last from 1 day to 2 weeks, after which bovine besnoitiosis progresses in two sequential phases: the febrile acute stage (anasarca) and the chronic stage (scleroderma) (21, 24).
The acute stage of bovine besnoitiosis, which can go undetected, starts with fever, when tachyzoites are invading the blood vessel endothelia. Vascular permeability then increases, producing edema. Degenerative and fibrinoid necrotic vascular lesions, vasculitis, and thrombosis also develop, mainly in skin and testes. Lymphadenopathy, inappetence, hyperemia of the skin, orchitis (see ), and lameness can occur. In severe cases, edema in pulmonary alveolar and interstitial tissues, accompanied by pneumonitis and emphysema, can also cause severe respiratory disorders.
Scrotum of a bull with orchitis due to besnoitiosis.
Courtesy of Dr. Gema Álvarez García.
Also during the acute phase, bulls can develop testicular degeneration, leading to sterility. This is driven by thermoregulation failure resulting from vascular lesions, which are often aggravated by tissue cysts occluding the blood vessel lumen and incipient scrotal skin lesions. Furthermore, damage to the blood-testis barrier induced by infiltrating immune cells might impair spermatogenesis during orchitis.
In lactating cows, an initial "milk drop syndrome" can occur during the acute phase of besnoitiosis; however, chronically infected cows typically recover, and milk production eventually returns to near-optimal levels.
Chronic besnoitiosis in cattle is produced by the slow replication of bradyzoites inside tissue cysts that have tropism for connective tissue—especially the superficial layers of skin, of the mucous membranes of the upper respiratory tract, and of the vestibulum vaginae (see )—and in bulls, in the testes and epididymis. Tissue cysts in the scleral conjunctiva are pathognomonic of chronic besnoitiosis (see ).
Pearls & Pitfalls
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Perineal region of a cow with besnoitiosis. Tissue cysts (arrow) are evident in the vestibulum vaginae.
Courtesy of Dr. Gema Álvarez García.
Eye of a cow with besnoitiosis. Cysts (arrow) are evident as pinpoint white-to-glistening papules on the scleral conjunctiva.
Courtesy of Dr. Gema Álvarez García.
Other typical clinical signs of chronic besnoitiosis in cattle are cutaneous lesions, such as hyperkeratosis and skin folding in the neck (see ), perineal region, scrotum (see ), and udder (see ); loss of necrotic epidermis; scars and nodules in the udder; and testicular atrophy and induration. During the chronic phase, scrotal skin lesions and cysts located in the epididymis (interstitial tissue and seminiferous tubules), as well as in the tunica dartos, vaginalis, and albuginea, also alter testicle cooling, resulting in degenerated seminiferous tubules and azoospermia.
Bull with besnoitiosis-related skin folding and hyperkeratosis, which are especially evident in the neck and proximal hindlegs.
Courtesy of Dr. Gema Álvarez García.
Scrotum of a bull with besnoitiosis. This bull has excessive folding of the scrotal skin.
Courtesy of Dr. Gema Álvarez García.
Cow with besnoitiosis. Note the hyperkeratosis in the udder (center arrows). Hyperkeratosis is also evident in the distal aspect of the limbs (upper right arrow).
Courtesy of Dr. Gema Álvarez García.
Death can occur in both the acute and chronic stages of besnoitiosis; however, the mortality rate is low. Severely affected bulls can become sterile during either acute or chronic infection. Affected cattle remain carriers for life.
The clinical signs of besnoitiosis in goats are similar to those in cattle. Infected donkeys also have similar clinical signs; however, the signs are often more severe than in horses.
Prevention and Control of Besnoitiosis
There are no effective treatments and no licensed vaccines for besnoitiosis in any species, with the exception of a live vaccine used in bulls in Israel. Strategies for control and prevention in herds of animals are, therefore, paramount and include the following:
biosecurity measures
early and accurate diagnosis to prevent transmission within the herd
serological testing to detect subclinical infections
selective culling of animals with clinical signs or lesions, low body condition scores, and poor productivity
separation of seropositive animals from the herd, with separate management
regular clinical and serological monitoring of the herd
regular monitoring for clinical signs and sperm quality before breeding season
use of insect repellents and ectoparasiticides indoors and outdoors; integrated control strategy for stable flies (Stomoxyscalcitrans)
Control strategies for besnoitiosis must be customized for individual herds on the basis of seroprevalence data, clinical impact, and the viability of implementing specific management measures. Management decisions should be based on both clinical inspection (to identify clinically affected animals) and serological analysis (to detect subclinically infected animals). The clinical veterinarian must assess the need for supportive care during the acute phase to manage fever or pain associated with potential lameness.
Biosecurity measures are key to controlling besnoitiosis. Risky practices, such as sharing pastures or bulls, should be avoided. New entries to the herd should be tested and diagnosed during the quarantine period to prevent disease introduction to the herd. Because iatrogenic transmission via syringes is possible, careful protocols should be followed. Use of repellents and ectoparasiticides is recommended, both indoors and outdoors.
Early and accurate diagnosis is essential to prevent and avoid transmission of besnoitiosis within the herd. The appearance of cysts in the scleral conjunctiva and vestibulum vaginae is a useful diagnostic indicator during clinical inspection, and clinical cases can be confirmed by skin biopsy and microscopic observation of tissue cysts. However, serological testing is essential for detecting subclinical infections.
In cattle herds, selective culling of animals with clinical signs or skin lesions, including tissue cysts in sclera or vestibulum vaginae, is recommended, as is the culling of unproductive cattle and cattle with low body condition scores. However, culling of seropositive animals or physical separation of infected animals from noninfected animals, in combination with appropriate reproductive management (noninfected cows bred with noninfected breeding bulls, and infected cows artificially inseminated or bred with seropositive fertile breeding bulls) is economically feasible in herds with low seroprevalence (< 10%).
In cattle herds, seropositive cattle should be progressively replaced by healthy, B besnoiti–seronegative cattle. Alternatively, PCR analysis of skin biopsy samples from the tail base might help identify potential subclinical seropositive superspreaders, as parasite transmission is dependent on parasite load. Although the proportion of superspreaders is generally low on farms exhibiting high seroprevalence, the rapid culling of superspreaders could yield good results in the medium term.
In addition to clinical monitoring, serological monitoring of the herd should be included in the regular herd health program. Regular monitoring should involve the following:
two sequential samples tested by immunoblot or validated ELISA
for replacement cattle > 6 months old: regular annual monitoring by immunoblot or validated ELISA
for breeding bulls: regular annual monitoring before breeding season by immunoblot or validated ELISA
for the whole herd: at least annual regular clinical inspection in Besnoitia-free herds in regions where the disease is endemic
Monitoring of sperm quality before breeding season (see Breeding Soundness Examination of Bulls) should also be a part of the regular herd health program.
Key Points
Besnoitiosis is characterized mainly by skin lesions during the chronic phase of disease; scleral conjunctival tissue cysts are pathognomonic of chronic besnoitiosis.
Besnoitiosis impairs fertility in cattle. Infected bulls can develop orchitis that results in sterility, during either acute or chronic infection.
Subclinically infected cattle are carriers. However, not all seropositive animals contribute equally to parasite transmission in the herd; that is dependent on the parasite load.
Serological testing, biosecurity measures, and adequate reproductive and selective culling strategies are key components of a besnoitiosis control program.
For More Information
Alvarez-García G, García-Lunar P, Gutiérrez-Expósito D, Shkap V, Ortega-Mora LM. Dynamics of Besnoitia besnoiti infection in cattle. Parasitology. 2014;141(11):1419-1435.
Gazzonis AL, Ferre I, Alvarez-Garcia G. Bovine besnoitiosis. In: Simões J, ed. Encyclopedia of Livestock Medicine for Large Animal and Poultry Production. Springer; 2025.
González-Barrio D, Diezma-Díaz C, Tabanera E, et al. Vascular wall injury and inflammation are key pathogenic mechanisms responsible for early testicular degeneration during acute besnoitiosis in bulls. Parasit Vectors. 2020;13(1):113.
Gutiérrez-Expósito D, Ferre I, Ortega-Mora LM, Álvarez-García G. Advances in the diagnosis of bovine besnoitiosis: current options and applications for control. Int J Parasitol. 2017;47(12):737-751.
Oryan A, Silver IA, Sadoughifar R. Caprine besnoitiosis: an emerging threat and its relationship to some other infections of ungulates by Besnoitia species. Res Vet Sci. 2014;97(1):1-7.
References
Schares G, Joeres M, Rachel F, et al. Molecular analysis suggests that Namibian cheetahs (Acinonyx jubatus) are definitive hosts of a so far undescribed Besnoitia species. Parasit Vectors. 2021;14(1):201. doi:10.1186/s13071-021-04697-3
Álvarez-García G. From the mainland to Ireland—bovine besnoitiosis and its spread in Europe. Vet Rec. 2016;178(24):605-607. doi:10.1136/vr.i3175
Talafha AQ, Al-Majali AM, Ababneh MM, Abutarbush SM. Epidemiologic study on Besnoitia besnoiti infection in dairy herds in Jordan. Parasitol Res. 2015;114(7):2491-2497. doi:10.1007/s00436-015-4448-5
Kula D, Gökpınar S. Seroprevalence of Neospora caninum and Besnoitia besnoiti in Cattle in Oğuzlar Region. Turkiye Parazitol Derg. 2021;45(2):108-112. doi:10.4274/tpd.galenos.2020.7075
Özdal N, Oğuz B, Orunç Kılınç Ö, Karakuş A, Değer S. Prevalence of ELISA-detected specific antibodies against Besnoitia besnoiti in cattle of the Eastern and Southeastern Anatolian regions, Turkey. Iran J Vet Res. 2019;20(2):143-146.
Ashmawy KI, Abu-Akkada SS. Evidence for bovine besnoitiosis in Egypt-first serosurvey of Besnoitia besnoiti in cattle and water buffalo (Bubalus bubalis) in Egypt. Trop Anim Health Prod. 2014;46(3):519-522. doi:10.1007/s11250-013-0522-0
Fereig RM, Salama DB, Salem FK, et al. Frequency of Besnoitia besnoiti and Neospora caninum antibodies in cattle and small ruminants from greater Cairo and Beni Suef governorates, Egypt. Vet Parasitol Reg Stud Reports. 2024;53:101078. doi:10.1016/j.vprsr.2024.101078
Kuraa HM, Youssef ZMA, Mahmoud FS, Malek SS. Seroprevalence of Besnoitia besnoiti in Assiut Governorate, Egypt. Open Vet J. 2022;12(5):754-761. doi:10.5455/OVJ.2022.v12.i5.21
Ai-Haboub T, Albarrak SM, Elsify A, Omar MA. First serological evidence and risk factor analysis of Neospora caninum and Besnoitia besnoiti infections in cattle and sheep from three regions of Saudi Arabia. Vet World. 2025;18(10):3229-3242. doi:10.14202/vetworld.2025.3229-3242
Al-Farwachi MI, Mohammad HA, Al-Robaiee IA. Prevalence of bovine besnoitiosis in Mosul city, Iraq. Iraqi J Vet Sci. 2025;39(3):639-645. doi:10.33899/IJVS.2025.157692.4140
Lee HS, Bak UB, Moon MH, Shin JU. Studies on bovine besnoitiosis in Korea: II. A survey on incidence in the enzootic region. Kisaengchunghak Chapchi [Korean J Parasitol]. 1970;8(3):76-80. doi:10.3347/kjp.1970.8.3.76
Gutiérrez-Expósito D, Ortega-Mora LM, Gajadhar AA, García-Lunar P, Dubey JP, Alvarez-García G. Serological evidence of Besnoitia spp. infection in Canadian wild ruminants and strong cross-reaction between Besnoitia besnoiti and Besnoitia tarandi. Vet Parasitol. 2012;190(1-2):19-28. doi:10.1016/j.vetpar.2012.06.017
Sambo SJ, Ibrahim ND, Esievo KA, Hambolu JO, Makoshi M., Tekdek LB. First incidence of caprine besnoitiosis in Kaduna State of Nigeria. J Anim Vet Adv. 2007;6:673-675.
Elsheikha HM, Schares G, Paraschou G, Sullivan R, Fox R. First record of besnoitiosis caused by Besnoitia bennetti in donkeys from the UK. Parasit Vectors. 2020;13(1):279. doi:10.1186/s13071-020-04145-8
Pols J, Alexander RA, Clark R, Louw JG, De Kock VE. Studies on bovine besnoitiosis with special reference to the aetiology. Onderstepoort J Vet Res. 1960;28(3):265.
Tinkler SH, Villa L, Manfredi MT, Walshe N, Jahns H. First report of Besnoitia bennetti in Irish donkeys: an emerging parasitic disease in Europe. Ir Vet J. 2024;77(1):2. doi:10.1186/s13620-024-00263-2
Ness SL, Peters-Kennedy J, Schares G, et al. Investigation of an outbreak of besnoitiosis in donkeys in northeastern Pennsylvania. J Am Vet Med Assoc. 2012;240(11):1329-1337. doi:10.2460/javma.240.11.1329
Olias P, Schade B, Mehlhorn H. Molecular pathology, taxonomy and epidemiology of Besnoitia species (Protozoa: Sarcocystidae). Infect Genet Evol. 2011;11(7):1564-1576. doi:10.1016/j.meegid.2011.08.006
Alvarez-García G, Frey CF, Mora LM, Schares G. A century of bovine besnoitiosis: an unknown disease re-emerging in Europe. Trends Parasitol. 2013;29(8):407-415. doi:10.1016/j.pt.2013.06.002
Gazzonis AL, Alvarez Garcia G, Maggioni A, et al. Serological dynamics and risk factors of Besnoitia besnoiti infection in breeding bulls from an endemically infected purebred beef herd. Parasitol Res. 2017;116(4):1383-1393. doi:10.1007/s00436-017-5418-x
Bigalke RD. New concepts on the epidemiological features of bovine besnoitiosis as determined by laboratory and field investigations. Onderstepoort J Vet Res. 1968;35:3-137.
Jacquiet P, Liénard E, Franc M. Bovine besnoitiosis: epidemiological and clinical aspects. Vet Parasitol. 2010;174(1-2):30-36. doi:10.1016/j.vetpar.2010.08.013
Diezma-Díaz C, Jiménez-Meléndez A, Fernández M, et al. Bovine chronic besnoitiosis in a calf: Characterization of a novel B. besnoiti isolate from an unusual case report. Vet Parasitol. 2017;247:10-18. doi:10.1016/j.vetpar.2017.09.017
Basson PA, McCully RM, Bigalke RD. Observations on the pathogenesis of bovine and antelope strains of Besnoitia besnoiti (Marotel, 1912) infection in cattle and rabbits. Onderstepoort J Vet Res. 1970;37:105-126.
