key: cord-0870215-egvf8hoq authors: Duszynski, Donald W.; Kvičerová, Jana; Seville, R. Scott title: Chapter 19 Species Inquirendae in the Carnivora date: 2018-12-31 journal: The Biology and Identification of the Coccidia (Apicomplexa) of Carnivores of the World DOI: 10.1016/b978-0-12-811349-3.00019-0 sha: 28394486c24c275b279ca6dbb6078d43ce946988 doc_id: 870215 cord_uid: egvf8hoq Abstract There are at least 483 reports of carnivore apicomplexans in the literature citing 17 genera or generic categoriesthat have not been described sufficiently or there was so little information provided by the author(s), that their validity, and sometimes even their identity is in question. These include: 2 “Apicomplexa protozoa,” 1 Besnoitia sp., 34 Coccidia-like or Coccidia spp., 135 Cryptosporidium spp., 2 Cyclospora spp., 26 Cystoisospora spp., 36 Eimeria spp., 3 Hammondia-like forms; 7 Hammondia–Neospora-like forms; 7 Hammondia–Toxoplasma-like forms; 14 Hepatozoon spp., 1 Hoareosporidium sp., 72 Isospora spp., 1 Klossia sp., 1 “oocysts,” 135 Sarcocystis spp., and 3 Toxoplasma-like forms. All are considered species inquirendae. In addition, two names, Isospora novocati of Pellérdy, 1974b and Isospora mustelae of Galli-Valerio, 1932, are considered nomen nuda, by definition. There is no record that 37/72 (51%) Caniformia genera, and all their 60 species, and 37/54 (68.5%) Feliformia genera, and all their 49 species, have any of these abbreviated identifications attached to them because most, if not all, likely were never surveyed for intestinal coccidian parasites. And, in the 52 genera that have been surveyed, the vast majority of their 177 species have not yet been looked at for coccidia. Surveys of various animal and plant species or populations are on-going events that occur worldwide on a regular basis by biologists of every persuasion (entomologists, marine biologists, botanists, mammalogists, ornithologists, herpetologists, etc.) . Historically, however, collecting parasites or even some parasite stages (e.g., surface organisms or fecal stages) have not been part of this survey equation, and parasitic faunas got ignored. However, in recent decades parasites have achieved recognition as important and abundant components of ecosystems, and whole organism surveys now often make at least cursory attempts to determine the presence and identification of some of the parasites of a particular host plant or animal species. In this vein, fecal material is often collected from captured hosts and later examined for parasite transmission stages (cysts, eggs, oocysts) that allow biologists to make either superficial identifications (to parasite genus) from distinctive propagules or in many cases to identify new morphological forms (from oocysts) that can result in new species descriptions. In the former instances, parasite stages may be identified quite superficially (an amoeba, a ciliate, a sporozoan) or they may be more specifically characterized (a Toxocara egg, an eimerian, an Isospora sp., a Sarcocystis sporocyst, etc.). It is these species "identifications" that, under certain circumstances, can provide useful information, but only in a general way; the forms with these kinds of names are listed in this chapter because no full specific identifications were made. We now know that at least four genera covered in our book (Besnoitia, Cystoisospora, Sarcocystis, Toxoplasma) produce developmental stages in hosts that may be prey items of carnivores. For example, some Sarcocystis species from omnivorous and/or herbivorous hosts that usually employ carnivores as definitive host (e.g., roe deer, Capreolus capreolus), but in which there is no mention of a possible carnivore host, will not be mentioned here because there is no way to connect them to any member of the Carnivora, even though the association may have strong circumstantial evidence (e.g., López et al., 2003) . The International Code of Zoological Nomenclature uses this designation for "a species of doubtful identity needing further investigation." Implicit in this definition for the coccidia is that the taxonomic "species" has been named in a published document, but without the existence of a "type specimen" of any kind (e.g., line drawing, photosyntype, stages in tissue sections, oocysts in preservative, etc.) and without quantitative and qualitative data on the most widely available stage in the life cycle, the sporulated oocyst, to distinguish it from other similar morphotypes or perhaps closely-related species. All the forms that we include in this chapter have lots of missing data that prevent them from having valid binomial designations. Remarks: Two neonatal male red panda littermates were submitted for necropsy. One animal was found dead with no prior signs of illness; the other had a brief history of labored breathing. Postmortem examination revealed disseminated protozoal infection. To characterize the causative agent, transmission electron microscopy (TEM), immunohistochemistry (IHC), polymerase chain reaction (PCR) and amplification, and nucleic acid sequencing were performed. IHC was negative for Toxoplasma gondii and Neospora caninum but was positive for Sarcocystis spp. TEM of cardiac muscle and lung revealed numerous intracellular apicomplexan protozoa within parasitophorous vacuoles (PV). PCR and nucleic acid sequencing of partial 18S rRNA and the internal transcribed spacer (ITS-1) region confirmed a Sarcocystis sp. that shared 99% sequence homology to S. neurona and S. dasypi. This was the first report of sarcocystosis in red pandas. Zoll et al. (2015) believed that their histopathological, immunohistochemical, molecular, and ultrastructural findings support vertical transmission resulting in fatal disseminated disease, but they did not assign a binomial name to the parasite of the red panda, so it must be considered a species inquirenda, at least for now. GENUS CANIS L., 1758 (6 SPECIES) Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Shelton et al. (1968) found what they described as subcutaneous nodules and a nonfebrile wasting illness in a 6-month-old border collie in Gallatin, Missouri, USA. Nodules were 10-12 mm wide and located on the lateral surface of the right maxilla. One lump was excised and prepared for standard H & E histological examination. Frequently, trophozoites, meronts with mature merozoites, and micro-and macrogametocytes were all discovered in the same host cell. Mature meronts were 11-20 in fixed tissue, and merozoites appeared to be arranged radially from a nuclear remnant of the meront. Merozoites in tissues were 5.5-6 × 2.0, crescent-shaped with one end thicker and blunter than the other, and a thick basophilic N was near the center of each merozoite. There were 16-20, sometimes 30, merozoites per meront. Microgametocytes were spheroidal, 8.5-11.5 wide. Spheroidal macrogametocytes, 8-9 wide, were the most prevalent structures seen, and each had a pale N with a distinct, centrally-located karyosome. Levine and Ivens (1981) thought these meronts appeared to be more like Besnoitia than any other genus, but they were unaware at the time that Caryospora bigenetica from rattlesnakes could produce similar infections in some mammals (see Chapter 4). Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Rinaldi et al. (2008) examined canine fecal samples in urban Naples, Campania region, southern Italy. Samples were examined by a commercial ELISA kit (ProSpecT ® Cryptosporidium Microplate Assay) for the presence of coproantigen of Cryptosporidium; 7/415 (2%) samples were positive, but the species/genotype was not identified, and because samples were collected from the ground, we will never know whether they were really canine or not. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: During their survey on gastrointestinal parasites in dogs and cats, Scorza and Lappin (2017) examined fecal samples of dogs on the Pine Ridge Indian Reservation, South Dakota, USA. Samples were examined by centrifugation-flotation, and by a commercial immunofluorescence assay for Cryptosporidium (Merifluor Crypto/Giardia kit). PCR amplification of the HSP70 gene was performed with immunofluorescence-positive samples. Cryptosporidium was detected in 6/84 (7%) samples, 2 of which could not be identified further (the other 4 were C. canis). Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Shukla et al. (2006) examined fecal samples of domestic dogs in the Niagara region, Ontario, Canada, by a fecal concentration method, acid-fast staining, and Cryptosporidium enzyme immunoassay (EIA) using the ProSpecT Cryptosporidum Microplate Assay. All samples were from dogs in one veterinary clinic; Cryptosporidium was detected in 5/68 (7%) samples, and all were found by the EIA test; neither of the other two methods detected Cryptosporidium. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Simonato et al. (2017) examined canine fecal samples from public green (270 samples) and urban (435 samples) areas, Padua municipality, Veneto region, northern Italy. Samples were examined by sedimentationflotation, followed by nested PCR amplification and subsequent sequencing of 18S rRNA gene. For some reason, samples concurrently positive for Giardia favored amplification of giardial DNA over the cryptosporidial DNA, so these samples also were examined by a touch-down real-time PCR of the COWP gene. Simonato et al. (2017) said that 12/705 (<2%) samples were positive for Cryptosporidium spp., 1 sample identified as C. canis, and the other 11 as "C. parvum species complex." However, because the samples were collected from the ground, one may never know whether they were really canine or not. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Sisk et al. (1984) described intestinal cryptosporidiosis in two pups in Georgia, USA. A 6-week-old mixed-breed female was "in a weak, semicomatose condition" and died within 12 hours. It was negative for parvovirus, adenovirus, and herpesvirus. Necropsy revealed mild hyperemia of the small intestinal serosa, mild interstitial pneumonia, and numerous "oval to spherical, dense basophilic bodies 3-4 μm in diameter" in the ileum, attached to the enterocytes, or free in the lumen of the small intestine. "Organisms morphologically consistent with Cryptosporidium," including meronts, merozoites, and trophozoites were observed in the ileum by TEM. The second pup that had suffered seizures was a 6-week-old, mixed-breed, from a cattle farm. It was negative for viral and bacterial agents, but dilated crypts with cellular debris were observed in the small and large intestines, and "sparse coccoid bodies 3-4 μm in diameter" were attached to the surface of enterocytes in the ileum. "Organisms morphologically consistent with Cryptosporidium," also were observed in the ileum by TEM, but the pup likely died of intoxication because 7.0 ppm of the pesticide toxaphene was detected in its liver. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Tangtrongsup et al. (2017) examined fecal samples of dogs in the Chiang Mai province, Thailand; 36 from the Small Animal Hospital, Chiang Mai University, 9 from private clinics, 15 from shelters, and 49 from breeders. Samples were examined by immunofluorescence assay (Merifluor Crypto/Giardia kit) and by PCR amplification of HSP70 and 18S rRNA genes. The overall prevalence of Cryptosporidiumpositive samples was 34/109 (31%), mostly in dogs <1-year-old; 14 dogs (13%) were positive by immunofluorescence, and 21 dogs (19%) by either/both of the PCR assays, and 18/109 (16.5%) were concurrently infected with G. duodenalis. Original host: Canis latrans Say, 1823, Coyote. Remarks: Thompson et al. (2009) examined coyotes in southern Alberta and Saskatchewan, Canada, by sucrose flotation, immunofluorescence of monoclonal antibodies specific for Cryptosporidium (Crypt-a-Glo TM ), and PCR amplification with subsequent sequencing and phylogenetic analyses of 18S rRNA and HSP70 genes. Cryptosporidium was detected in 8/70 (11%) samples, all collected in winter, whereas no samples collected in summer were positive. This may be because in winter the coyotes were in poor condition and nutritionally compromised; 6/8 (75%) positive samples could not be successfully amplified by PCR. and Salem (2015) surveyed fecal samples from military (40 samples), nomadic (30 samples), and household dogs (60 samples) in Sharkia and Qalyubia provinces, Egypt. Samples were examined by centrifugation-flotation. Awadallah and Salem (2015) reported finding some C. cayetanensis oocysts in 1/130 (<1%) samples from a nomadic dog. Their photomicrograph is blurred, and the oocyst shown does not resemble a Cyclospora oocyst (in our opinion). Knowing that dogs are coprophagous, we suspect this is a spurious finding of an oocyst passing through the dog's gut after ingesting infected human feces. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Yai et al. (1997) reported an infection with Cyclospora sp. in each of two dogs from São Paulo, Brazil, 1 Siberian husky and 1 Rottweiler. Both dogs had a history or watery diarrhea, weight loss, and lethargy. However, their photomicrographs of (presumably) sporulated oocysts of Cyclospora were not convincing. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Palmer et al. (2008a) surveyed fecal samples of dogs from both urban and rural areas across Australia, which were collected from 59 veterinary clinics (810 samples) and 26 refuges (590 samples). Samples were examined by centrifugation-flotation. "I. (=C.) ohioensis complex" was detected in 49/1,400 (3.5%) dogs, mostly from refuges. Remarks: Gennari et al. (1999) examined fecal samples of domestic dogs from different areas of São Paulo, Brazil. Samples were examined by flotation and oocysts of Cystoisospora sp. were detected in 9/353 (2.5%) samples. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Gill et al. (1978) found unsporulated Isospora oocysts in the feces of dogs that were fed diaphragm from water buffalo (Bubalus bubalis) naturally-infected with macroscopic sarcocysts of S. fusiformis (but they did not examine the diaphragm for microscopic cysts). Four dogs that had never been fed meat each were fed 25 g of buffalo diaphragm, and all four dogs started shedding unsporulated oocysts on 9 (three dogs) or 10 DPI (one dog), and the infected dogs continued shedding oocysts daily for 15, 18, 23, and 25 days. Sporulation was completed in 8-16 hours at room temperature. Sporulated oocysts were L × W (n = 25): 18.2 × 13.3 (17.5-24 × 16-19) , L/W ratio: 1.1 (1.0-1.3), and M, OR, PG: all absent. Sporocysts were L × W (n = 25): 13.3 × 9.8 (11-16 × 9-11), L/W ratio: 1.4 (1.1-1.6), and SB, SSB, PSB: all absent, but with a large SR. Gill et al. (1978) said that these oocysts either were Hammondia or Isospora but neglected to assign their form to either. Levine and Ivens (1981, pp. 49-50) suggested this may be a Toxoplasma species, even though the sporulated oocysts are about the same size as those of C. burrowsi and C. ohioensis, but gave no reason to support calling it Toxoplasma sp. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Huber et al. (2005) surveyed fecal samples of 94 stray dogs from an animal shelter and 72 household pet dogs from the West Zone of Rio de Janeiro, Brazil. All the dogs were clinically healthy. Oocysts of Cystoisospora were detected in "some samples"; however, the number of positive samples was not given. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Yu et al. (2017) examined fecal samples of pet dogs with diarrhea from various regions in Beijing, China. Samples were examined by LM of wet fecal smears, and Cystoisospora oocysts were detected in 21/485 (4%) samples, mainly in poodles and golden retrievers, but the parasites were not further identified to the species. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Rao and Bhatavdekar (1957) described ovoidal to ellipsoidal oocysts from a dog in Bombay, India, which were L × W (n = 50): 26.8 × 19.8 (22-29 × 18-22) ; L/W ratio: 1.35. The oocysts had a MC and a M, the former was 5.8 (4-7) and the latter was 1.7 (1-4) wide. Unfortunately, the oocysts were unsporulated so sporocyst numbers and size were not recorded. Levine and Ivens (1981) believe these oocysts resemble those of E. arloingi from goats merely passing through the gut of the dog. Whatever they were, they were not a parasite of the dog and thus can only be relegated to species inquirenda. Remarks: Fok et al. (2001) examined fecal samples of dogs in eastern and northern Hungary and reported a presence of Eimeria spp., which they called "spurious parasites" (Table 1 , p. 97), in 18/490 (4%) dogs from a country town and a village, but gave no details. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Streitel and Dubey (1976) surveyed feces from stray dogs in a humane shelter in Ohio, USA and reported 2/500 (0.5%) samples to have oocysts of an Eimeria species. After sporulating the oocysts, they were inoculated via stomach tube, into one coccidia-free, 1-month-old lab-reared pup. The feces of the pup were monitored for 30 DPI, but no additional oocysts were ever found. Although there are several Eimeria species described from dogs, they suggested that, "Eimeria are probably accidental 'parasites' of dogs because the Eimeria sp. found in this survey was not infectious to a dog." Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Guillebeau (1916) described a small coccidium that he identified as E. stiedae in the liver cells of dogs, even though the oocysts only measured L × W: 12 × 7, far too small to be those of E. stiedai found in the liver of rabbits. Coccidial developmental stages in the liver of dogs would be an unusual finding, indeed, given all of the domestic dogs that have been examined worldwide in the last 100 years! Guillebeau's (1916) figures do not help in deciding even the nature of the organism at which he was looking; thus, another species inquirenda. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Cirak and Bauer (2004) examined fecal samples of healthy dogs from animal shelters in Germany to compare conventional coproscopical methods versus a commercial coproantigen ELISA kit, to best detect Cryptosporidium. Hammondia-like oocysts were found in 2/270 (<1%) dogs using the ZnCl 2 -NaCl flotation method. reserves (four in the North-West province, one in Limpopo province), South Africa. Specimens were screened for Babesia, Theileria, Hepatozoon, and Ehrlichia/Anaplasma species using PCR and reverse line blot assays. Two dogs were positive for Hepatozoon sp. Original hosts: Canis adustus Sundevall, 1878, Side-striped Jackal; and Lions, Cheetahs, Hyaenas, and Leopards. Remarks: McCully et al. (1975) studied hepatozoonosis in hyaenas, lions, jackals, cheetahs, and one leopard in the Kruger National Park, to compare possible symptoms with those seen in some dogs in South Africa. Meronts of Hepatozoon were found in many wild carnivores and they illustrated the progressive development of microschizonts. They reported meronts in the lung, myocardium, and skeletal muscle, and sometimes also in the spleen, liver, and lymph nodes. Gametocytes were present in leukocytes, but they saw very little of a host response to the presence of Hepatozoon infection, and there was no attempt to determine the species. Sporogony in ticks was reported in Rhipicephalus simus females removed from an infected hyaena and R. sanguineus adults fed on an infected jackal in the nymphal stage. Attempts to transmit Hepatozoon from a jackal to domestic dogs by means of ticks gave "inconclusive results." Original host: Canis lupus (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Pande et al. (1972) reported an unusual coccidium in the rectal contents of four pups slaughtered in India. It was similar to Cryptosporidium because of its asporous, tetrazoic oocysts, which they saw in fresh scrapings of the jejunum; however, it was intracellular in epithelial and subepithelial cells of the villi, rather than in the intracellular, but extracytoplasmic, location typical for Cryptosporidium species, so they decided to create a new genus for their organism. They measured their "oocysts" both extracellularly and intracellularly. Fresh "oocysts" were L × W (n = 50): 14.8 × 9.1 (12-17 × 8.5-10.5), L/W ratio: 1.6 (1.2-1.8); the OR was a prominent mass of large-sized granules, and sausage-shaped SZ were 9.3 × 2.7 (9-10.5 × 2.6-3). Intracellular "oocysts" were L × W (n = 25): 14.2 × 8.8 (12-17 × 7-10); L/W ratio: 1.6, and SZ were 8.2 × 2.4 (6-10.5 × 2-2.5). They said, "though the dimensions given for the oocysts in V. vulpis Wetzel, 1938 (from common red fox) are well within the range encountered in our material, the oocysts in this species, according to Pellérdy, could possibly be the freshly-shed sporocysts of some known or as yet unknown isosporan species." We are convinced that what Pande et al. (1972) reported actually were sporocysts of some Sarcocystis species developing in the dog's jejunal epithelial cells, which they mistook for oocysts of their new Hoareosporidium pellerdyi. We must conclude that this form can only be regarded as a species inquirenda. Synonyms: Cystospermium villorum intestinalium canis Rivolta, 1878, nomen nudum; Coccidium rivolta Grassi, 1879, pro parte; Coccidium rivoltae Leuckart, 1886, pro parte; Coccidium bigemina Stiles, 1891, pro parte; Coccidium bigeminum var. canis Railliet and Lucet, 1891; Diplospora bigemina Martin, 1909; Coccidium bigeminum Wigdor, 1918, pro parte; Isospora cati Marotel, 1922, pro parte; Lucetina bigemina (Stiles, 1891) Henry and Leblois, 1926; Isospora bigemina Gousseff, 1933, lapsus; Isospora bigemina var. bahiensis de Moura Costa, 1956; and others. Type host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Stiles (1891) found some parasites developing in the lamina propria of the domestic dog and called it Coccidium bigemina; Lindsay et al. (1997) suggested that he was likely looking at a Sarcocystis species because oocysts and sporocysts were seen to develop in the gut tissue and we agree. This organism was placed in the genus Isospora by Lühe (1906) and 2 decades later, Wenyon (1926a) suggested there were two "races" of I. bigemina, a large and a small race. The larger race, which was excreted as sporocysts and sporulated oocysts was a Sarcocystis species, and the smaller race, excreted as unsporulated oocysts, is now known to be Hammondia heydorni, an obligatory heteroxenous parasite (Heydorn et al., 1975c; Lindsay et al., 1997) . Nukerbaeva and Svanbaev (1974, 1977) collected oocysts from domestic dogs, which they identified as I. bigemina. They tried to transmit their oocysts to one 40-day-old Arctic fox (V. lagopus), and two 1-year-old minks (N. vison), but none of these animals became infected. They were successful in transmitting their oocysts to a control dog, which shed oocysts in its feces on the 8th DPI. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: In Fort Collins, Colorado, USA, Gassner (1940) examined dogs and said that 253/320 (79%) dogs were infected with the small form of Isospora bigemina and discharged either sporocysts or oocysts in their feces; Because free sporocysts and sporulated oocysts were being discharged in the feces, we now know Gassner (1940) was seeing a species of Sarcocystis. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Cirak and Bauer (2004) examined fecal samples of healthy dogs with normal stools in central Germany; oocysts of Isospora burrowsi/Isospora ohioensis were found in 3/270 (1%) dogs. They were unable to determine identity beyond that general observation. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Fontanarrosa et al. (2006) surveyed fecal samples of dogs in southern Buenos Aires, Argentina. They detected an "Isospora ohioensis complex" of oocysts in 263/2,193 (12%) samples, mostly in young dogs 0-6-months-old. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Blazius et al. (2005) examined fecal samples of stray dogs in Itapema City, Santa Catarina, Brazil, and found Isospora oocysts they did not identify to species in 10/158 (6%) samples. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Causapé et al. (1996) Original host: Canis lupus (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Collins et al. (1983) reported Isospora spp. in 6/110 (5.5%) fecal samples from dogs in Sydney, Australia, which were examined for "sporozoa." No other information was given. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: de Oliveira et al. (1990) examined fecal samples of 11,563 dogs from Uberlȃndia, Minas Gerais, Brazil and found Isospora oocysts in 148/3,202 (5%) samples but did not identify them to species. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Dubey et al. (1978b) found a coccidium in the villar epithelium, lamina propria, and intestinal glands in the distal half of the ileum, cecum, and colon of a 10-week-old puppy that apparently died from the infection in Ohio, USA. They said that sporulation was exogenous and oocysts were L × W: 19 × 16 (16-23 × 14-20) in fecal smears and 13 × 11.5 (12-17 × 10-13) in tissue sections and speculated that the infection might be a mixture of I. neorivolta and I. ohioensis, but had to conclude that they could not identify the parasite because the culture was "inadvertently discarded." They also reported at least two structurally different meronts and three different-sized merozoites in tissue stages of the pup. For more descriptive information on these various stages, see either Dubey et al. (1978b) or Levine and Ivens (1981, p. 22 Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Jaskoski et al. (1982) reported collecting fecal samples from public areas (streets, parks, etc.) in Edgewater and Rogers Park, Chicago, Illinois, USA, areas where pet dogs were kept inside and leashed when outside. In 1970, 14/846 (<2%) samples had Isospora oocysts, but during 1979-1980, only 1/806 (0.1%) samples was found with Isospora-type oocysts. Unfortunately, fecal samples were not collected directly from dogs, so whether they all were canine was highly probable, but unknown for certain. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Jordan et al. (1993) Original host: Canis lupus L., 1758, Wolf. Remarks: Mech and Kurtz (1999) found three 4-month-old C. lupus pups in the Superior National Forest of Minnesota, USA, which died in August/September, 1997, apparently of coccidiosis. Two of the pups had hemorrhagic feces and the third had a severely autolyzed intestine. The intestinal mucosa of two pups had many developmental stages in both enterocytes and in the lamina propria, which the authors attributed to an Isospora (probably Cystoisospora) species. No other information was provided. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Meloni et al. (1993) Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Miller et al. (2003) reported a concurrent infection with Cryptosporidium canis and Isospora spp. in an 8-week-old female Yorkshire terrier puppy in Tifton, Georgia, USA. The puppy suffered weakness and diarrhea. Histology revealed severe gastric and intestinal cryptosporidiosis, severe intestinal isosporiasis, and thymic atrophy with lymphoid depletion. Numerous sexual and asexual stages of Isospora spp. were observed in the luminal and crypt epithelial cells of the small intestine. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Nieschulz (1925) discussed Isospora infections in dogs and cats in the Netherlands; in addition to brief descriptions of I. (=C.) rivolta and I. (=C.) felis, he said he saw "free sporocysts (not whole oocysts) of an Isospora species in intestinal material of a young dog," which were ellipsoidal, L × W: 15.5 × 9.2 (14-17 × 8-10). Clearly, he saw and measured sporocysts of a Sarcocystis species. Original host: Canis aureus L., 1758, Golden Jackal. Remarks: Yakimoff and Lewkowitsch (1932) reported finding oocysts in one golden jackal in Azerbaijan (former USSR). Their oocysts were spheroidal or slightly ovoidal, L × W: 21.2 × 17.1-18.0, and had ellipsoidal sporocysts that were 16 × 11 (sic) (13-16 × 9-11); oocysts lacked a M, OR, and PG, and sporocysts lacked SB, SSB, PSB but were illustrated to have a granular SR. They said that sporocysts were found free in the intestine, which suggests they had seen a Sarcocystis sp., and that during transportation to Leningrad almost all of the oocysts' walls broke and disappeared, which would support this idea. Yakimoff and Lewkowitsch (1932) failed to transmit this form to the domestic dog with sporulated oocysts. Glebezdin (1978) surveyed wild mammals in southwestern Turkmenistan from 1974 to 1977; he reported finding, in C. aureus, ellipsoidal oocysts of I. theileri that measured L × W: 23.0 × 17.5 (22-25 × 17-20) , L/W ratio: 1.3; sporocysts were L × W: 11.6 × 9.1 (11-14 × 8-11). Levine and Ivens (1981, p. 51) hoped that someone will find this form again and suggested, "it too should be restudied if it is ever rediscovered." The data available are so sketchy and tentative, that we strongly agree. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Carlslake et al. (2017) examined dog fecal samples in Samoa, South Pacific. All dogs were mixed-breed and free-roaming, and Carlslake et al. (2017) reported the presence of "oocysts" in 9/204 (4%) dogs. Synonym: Sarcocystis capreolicanis Erber, Boch, and Barth, 1978. Definitive hosts: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog; Vulpes vulpes (L., 1758), Red Fox. Intermediate host: Capreolus capreolus (L., 1758), European Roe Deer. Remarks: The relationship of this species to S. gracilis and S. sibirica is unknown, but we can find no other information on this species that was listed as a valid species by Levine (1986) . Levine and Ivens (1981) did not mention this species nor did Dubey et al. (2015a) . Sedlaczek and Wesemeier (1995) examined 66 roe deer in Germany and Poland to compare the ultrastructure of their sarcocysts. They identified three distinct species, and when defining one of them, S. capreolicanis Erber et al. (1978) , they said that Levchenko's (1963) "species" is not recognizable in its original description and, therefore, should be a species inquirenda. We agree. Definitive hosts: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog; Felis catus L., 1758, Domestic Cat. Intermediate host: Bos taurus L., 1758, Aurochs. Remarks: Mehlhorn et al. (1975b) published a paper in which they tried to compare the ultrastructure of sarcocysts of (what they called) S. fusiformis in the muscles of naturally-infected cows with sarcocysts in cow muscle after experimental infection by sporocysts of I. hominis and sporocysts of the large forms of I. bigemina from the dog and cat. In naturally-infected cows they found two kinds of sarcocysts, probably S. fusiformis from cats and S. cruzi from dogs, but in their defense, these life cycles were still unknown at the time. They concluded that one type of sarcocyst in cows was produced by the large form of I. bigemina from dogs and the second type of sarcocysts "were caused by infection with I. hominis and the large form of I. bigemina from the cat." Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Farmer et al. (1978) surveyed feces from 123 working sheepdogs in Gwynedd, 33 greyhounds from the London area, and 41 red foxes killed on Anglesey or in the Bangor area of England. They found sporocysts of Sarcocystis spp. in 35/123 (28%) sheepdogs, 8/33 (24%) greyhounds, and 7/41 (17%) red foxes (their Table 1 , p. 79). Based only on sporocyst dimensions, they said that S. porcifelis (syn. their S. miescheriana) sporocysts were identified in 11/47 (23%) sheepdogs and 2/8 (25%) greyhound samples they measured (their Table 3 , p. 79). At that early stage in time, when we knew little about the true identity and life cycles of most Sarcocystis species, using only sporocyst measurements to place them into the correct species diagnosis was a bit risky; and we now know that S. porcifelis (syn. their S. miescheriana) seems to only be transmitted by felids so we must relegate their form to a species inquirenda. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Capra hircus L., 1758, Goat. Remarks: Barham et al. (2005) surveyed goats slaughtered in the winter in northern Iraq and found three morphologically distinct types of sarcocysts, macrocysts (both fat and thin), and microcysts; macrocysts occurred in 281/826 (34%) goats, but these were not identified. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: During a coprological survey of dogs and cats in Germany, Barutzki and Schaper (2003) examined feces from both hosts in Freiburg, Germany; sarcocysts of Sarcocystis spp. were detected in 759/8,438 (9%) samples. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Blagburn et al. (1989) found a single sarcocyst in the biceps femoris muscle of a dog in Alabama, USA. The sarcocyst was spheroidal to ovoidal and measured 52 × 47 with a wall thickness of ∼2.3. Sarcocyst wall projections were barely visible with the LM, but under TEM they appeared as irregularly-spaced electrondense projections that were 1.5 × 0.9. Septa were visible in the sarcocyst and contained numerous, irregularly arranged bradyzoites, but no metrocytes were seen. The sarcocyst did not elicit an inflammatory response in the adjacent muscle tissue. Remarks: Dubey (1980a) fed 1 kg of tissues from bison, cattle, elk, goats, moose, and sheep to different Sarcocystis-free coyotes, and all 12 coyotes shed sporocysts of Sarcocystis in their feces with prepatent periods of 9-15 DPI. Dubey (1980a,b) said that sporocysts in coyote feces fed infected musculature of cattle, sheep, goats, and elk were all structurally similar and that this was the first report of the completion of the life cycle of Sarcocystis species in moose and bison. One other cross-transmission experiment indicated that one goat Sarcocystis species completed its life cycle in both the dog and coyote and that ovine Sarcocystis is not transmissible to goats. None of the Sarcocystis species were identified, thus all must remain species inquirendae. Remarks: Dubey et al. (1991a) examined a litter of eight dogs born to a primiparous 2-year-old Rottweiler in Maryland, USA. All of the pups had medical issues, but one (dog 1), at 7-weeksold, was listless and anorectic with mild anemia after surgery for entropion. It was killed 2 days later and specimens of liver and small intestine were examined histologically. Protozoan meronts were seen free in the cytoplasm of hepatocytes adjacent to necrotic foci; the parasites were without a PV, divided by merogony, and resembled Sarcocystis structurally and antigenically, as it reacted with Sarcocystis antiserum. It was uncertain how the pup became infected, but was noteworthy to find Sarcocystis in the visceral tissues of dogs. There was no way to know what species they saw so this must be considered as another species inquirenda. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Dubey et al. (1992a) examined paraffin-embedded tissue samples of liver, spleen, lungs, and kidneys of a 2-day-old female bullmastiff puppy that died 2½ days after birth. Initially deparaffinized tissues reacted with antisera to T. gondii, N. caninum, and S. cruzi. To better identify the protozoan, they studied tissue sections for developmental stages, documented some of the pathology, and isolated meronts to find whether they reacted with S. cruzi antiserum, but not with antisera to T. gondii or N. caninum. Lesions were seen in all tissues examined. The main lung lesion was interstitial pneumonia characterized by infiltration of mononuclear cells and neutrophils in the alveolar wall and alveolar hemorrhage and necrosis due to numerous meronts in the vascular endothelium. Hepatic lesions included necrosis of hepatocytes, moderate perivascular infiltration of mononuclear cells, and the presence of meronts and merozoites in hepatocytes and Kupffer cells. Renal lesions showed necrosis of glomeruli associated with all developmental stages of meronts, and multifocal areas of necrosis were present. Mature meronts with merozoites were L × W (n = 13): 14-20 × 10-20, contained 12-28 merozoites, and they are divided by endopolygeny. They were located in the host cell cytoplasm without a PV. The authors said, "The parasite was antigenically and structurally identical to the newly named protozoan Sarcocystis canis from Rottweiler dogs (Dubey and Speer, 1991)," but they declined to say that Sarcocystis found in this Louisiana puppy was S. canis. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Dubey et al. (1991c) examined a 10-month-old Rottweiler from Illinois, USA that had multiple cutaneous abscesses over its body, particularly in its hind limbs that were swollen, hot, and painful. The dog died shortly after a few biopsies were taken, so portions of skin, liver, lungs, kidney, and lymph nodes were processed for examination. Toxoplasma gondii, N. caninum, and Caryospora species are parasites that can cause dermatitis and disseminated infection in dogs, but these were all eliminated by not reacting with the appropriate antisera and/or physical structure of the protozoan stages found. Dubey et al. (1991c) concluded, "the present case resembles the Sarcocystis parasite that causes fatal encephalomyelitis in horses, cattle, and sheep. Although the central nervous system in the infected dog was not examined, its littermate died of protozoan encephalomyelitis." Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate hosts: Ovis aries L., 1758, Red Sheep; Bos taurus L., 1758, Aurochs. Remarks: Dubey and Streitel (1976c) did crossinfections between dogs, cats, sheep, pigs, and cattle from Iowa or Ohio, USA. Two dogs fed 100 sheep esophagi and hearts from Iowa shed sporocysts, but cats did not. Sporocysts were L × W (n = 11): 14.0 × 9.2 (13-15 × 9-10), L/W ratio: 1.5 and were shed for >8 days. Dogs shed large numbers of sporocysts and sporulated oocysts for >8 days after ingesting bovine tissue; sporocysts were L × W (n = 27): 15.0 × 9.5 (14-17 × 8-10), L/W ratio: 1.6. Because cats also shed sporocysts when fed bovine tissues, Dubey and Streitel (1976c) suggested that separate species of Sarcocystis parasitize cattle in different areas of the United States, but they did not suggest a name for any of these Sarcocystis species. Original host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Dubná et al. (2007) reported on a survey of canine and ground-collected fecal samples from various areas in and around Prague, and from rural areas in central Bohemia, Czech Republic. Sporocysts of Sarcocystis spp. were detected in 24/3,780 (<1%) samples. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Capreolus capreolus (L., 1758), European Roe Deer. Remarks: Entzeroth (1982b) studied the ultrastructure of gamonts and gametes and fertilization of a Sarcocystis species in dogs after they were fed sarcocysts from roe deer in Germany. Unfortunately, he did not attempt to name the species. He may have been dealing with the same species seen by Erber (1978) 4 years earlier (below) and both may have been working with S. gracilis, which is now known from the European roe deer. Nonetheless, we can only call it a species inquirenda based on the information provided by Entzeroth (1982b). Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Capreolus capreolus (L., 1758), European Roe Deer. Remarks: In Germany, Entzeroth et al. (1978) fed muscles of roe deer infected with sarcocysts to a coccidia-free fox, dog, and cat (species not stated). After a prepatency of 8 DPI, the fox shed sporulated sporocysts in its feces that were L × W: 14.5 × 8.5, and after the 11th day, unsporulated oocysts discharged by the fox were 13.1 × 11.6 and "resembled the small form of Isospora bigemina (Hammondia)." The dog shed sporulated sporocysts that were L × W: 15.6 × 10, with a prepatent period of 10-14 days, and sporocysts continued to be discharged for 51 days. The cat fed sarcocystinfected deer meat shed unsporulated oocysts on the 8th DPI that were L × W: 12.2 × 10.9. The small isosporan oocysts from the fox and the cat were inoculated into mice; Entzeroth et al. (1978) reported they found typical Toxoplasma cysts in the brain of the mice and cysts typical of Hammondia hammondi in the muscle of the mice with oocysts from the cat but that "oocysts from the fox did not cause any visible infection in mice." When they (Entzeroth et al., 1978) examined the muscle sarcocysts in roe deer with the TEM, they said there were three types of sarcosporidian cysts, but nothing else. Thus, more species inquirendae. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Odocoileus virginianus (Zimmermann, 1780), White-tailed Deer. Remarks: Entzeroth et al. (1982b) found 14/48 (29%) white-tailed deer infected with Sarcocystis, with a preference in location for the tongue. This was only about 4 months after Crum et al. (1981) named a species with sarcocysts in white-tailed deer tissue (especially tongue), and dogs as definitive hosts that discharged sporocysts as S. odocoileocanis. Entzeroth et al. (1982b) said their sarcocysts were spindle-shaped and measured 300-620 × 60-120, had a thin wall, and were divided into compartments by septa. They fed a lab-reared dog infected venison flesh and on 14 DPI the dog began to discharge sporulated sporocysts that were L × W: 14.9 × 10.6 (13.5-16.5 × 9-11); patency lasted at least 10 days, after which no fecal samples were checked. This deer/dog cycle is remarkably similar to that described by Crum et al. (1981) , and their sporocyst size, prepatent and patent periods also overlap, but the sarcocysts in the Michigan deer were larger than those reported by Crum et al. (1981) in deer from several southeastern states (e.g., 264 × 40 (150-536 × 30-51) versus 300-620 × 60-120). Unfortunately, Entzeroth et al. (1982b) were unaware of the paper published by Crum et al. (1981) and they chose not to name their species "until additional details of the life cycle are known." Thus, their form can only be considered a species inquirenda. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Epe et al. (1993) , in Germany, surveyed 3,329 fecal samples from dogs between 1984 and 1991 and found 100 (3%) to pass Sarcocystis spp. sporocysts, which were not identified to species. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Capreolus capreolus (L., 1758), European Roe Deer. Remarks: Erber (1978) reported three types of sarcocysts in the tongues and abdominal musculature of 391/421 (93%) C. capreolus, in West Germany. He fed raw muscles infected with his type 1 and type 2 sarcocysts to dogs, foxes, and cats. On 10-11 DPI, the dogs and foxes shed sporocysts in their feces that were L × W: 16-18 × 9-12, and the patent period lasted 50 days. The cats did not shed oocysts (Levine and Ivens, 1981) . To our knowledge, these forms were never studied again. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Hill et al. (1988) saw sarcocysts in the myocardium of a 2-year-old, spayed, female Doberman pinscher in Georgia, USA and said the dog's sarcocysts were similar in size and structure to sarcocysts they found in two cats, when examined by LM and TEM. The minor difference in sarcocysts from the dog (vs. cat sarcocysts) were a thinner layer of ground substance associated with the cyst wall and slightly larger bradyzoites. The cyst wall of both the dog and cats had striations, septa for compartmentalization, and fairly large bradyzoites. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Nanger granti (Brooke, 1872) (syn. Gazella granti), Grant's Gazelle. Remarks: Janitschke et al. (1976) found sarcocysts in Grant's gazelle in Tanzania, fed infected flesh to both cats and dogs, and reported finding sporocysts and sporulated oocysts in the feces of both. Sporocysts shed by the dog were L × W: 16 × 11 (13-18 × 8-12), L/W ratio, 1.45; the prepatent period in the dog was 10 days, and patency lasted 42 days. They thought they were dealing with two Sarcocystis species, but without further infection and/or molecular studies, it is not possible to determine which-was-which. Therefore, these must remain species inquirendae until they can be differentiated and named. There are at least three Sarcocystis species that have been named from gazelles, but only from sarcocysts in their muscle tissues. All three, S. gazellae Balfour, 1913 , S. mongolica Matschoulsky, 1947 , and S. woodhousei Dogel, 1916 , are mentioned in Levine (1986 and in Dubey et al. (2015a), but the carnivore definitive host is not known for any of them. Clearly a lot of work still needs to be done in sorting out the various Sarcocystis species in gazelles. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Katagiri and Oliveira-Sequeira (2008) examined fecal samples of 129 stray and 125 pet dogs from urban Botucatu, São Paulo State, Brazil. Samples were examined by multiple techniques. Sarcocystis sporocysts were found in 7/254 (3%) samples, 4 from strays and 3 from household dogs. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: King et al. (2012) mentioned briefly in the discussion of their paper on the presence of N. caninum in Australian Aboriginal dogs that they had seen "Sarcocystis spp. sporocysts in many of the dog feces examined," but gave no other information. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Phasianus colchicus L., 1758, Common or Ring-necked Pheasant. Remarks: Pecka (1990) found 3/90 (3%) pheasants, from two pheasant farms in the Czech Republic, infected with a Sarcocystis species with banana-shaped cystozoites, 6 × 2, and 46/90 (51%) were infected with a Sarcocystis species with lancet-shaped cystozoites, 14-16 × 2-3. A dog was infected experimentally with the latter species and pheasants were successfully reinfected, presumably with sporocysts from the dog, but this was not stated. Pecka (1990) said he suspected foxes to be the main final host for the pheasant species. No other information was given. Definitive host: Unknown. Intermediate host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Remarks: Sahasrabudhe and Shah (1966) reported finding sarcocysts in the muscles of an esophageal nodule of a dog in India. These sarcocysts had no septa, were 110-250 wide, and contained thousands of crescent-shaped merozoites, 4-5 × 1.5. They said that theirs was the first report of a Sarcocystis species from a domestic carnivore. Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. Remarks: Salb et al. (2008) examined fresh fecal samples of dogs presented to veterinary clinics from two remote northern Canadian communities (Fort Chipewyan, Alberta, Fort Resolution, Northwest Territories). The authors detected Sarcocystis sporocysts, but the number of positive samples and further details were not provided. Remarks: Soriano et al. (2010) surveyed dog fecal samples in urban (646 samples) and rural (1,298 samples) areas of Neuquén province, Patagonia, Argentina. Sarcocystis sporocysts were found in 110/1,944 (6%) samples, mostly from rural areas (99 of 110 samples). Definitive host: Canis lupus familiaris (syn. C. familiaris) L., 1758, Domestic Dog. Intermediate host: Unknown. (1976) surveyed feces from stray dogs in a humane shelter in Ohio, USA and found only 9/500 (2%) samples to have sporocysts of a Sarcocystis species. The sporocysts measured L × W: 13.5-16.2 × 8-11, but the species was neither identified nor was there an attempt to transmit it to another animal; thus, another species inquirenda. Gray, 1843) , Yak. Remarks: Wei et al. (1985) named two species of Sarcocystis from yaks, using LM, TEM, and SEM to distinguish between their sarcocysts, S. peophagi and S. poephagicanis. Wei et al. (1989) then examined 23 3-5-month-old fetuses taken from slaughtered yaks in Gan Su province, China and examined their tissues (heart, liver, kidney, mesenteric lymph nodes, femoral and diaphragm muscles, intestines) histologically for sarcocysts. They detected sarcocysts in 6/23 (26%) fetuses, three in diaphragm and three in myocardium and found meronts, but not clear merozoites. They did not name the species of Sarcocystis, even though they had named the only two known species in yaks (Wei et al., 1985) and did cross-transmission work with both species a year later (Wei et al., 1990) . This was not the first time that various stages of sarcocyst development were seen in fetal tissue; earlier, Munday and Black (1976) had reported Sarcocystis spp. in brain tissue of two aborted bovine fetuses, and Hong et al. (1982) found the vascular epithelium over all body parts contained immature and mature meronts of Sarcocystis in aborted bovine fetuses. These two studies, and their own work (Wei et al., 1985) , led them to conclude that at least some Sarcocystis species can be transmitted vertically from naturally-infected mothers to their fetuses. This is certainly an area that deserves further exploration. Definitive host: Unknown. Intermediate host: Canis mesomelas Schreber, 1775, Black-backed Jackal. Remarks: Wesemeier et al. (1995) examined fixed tissue sections of pieces of tongue from 25 black-backed jackals from Namibia in 1993. In one tongue (4%), using LM and TEM, they found two structurally different sarcocysts. Type 1 sarcocysts had a thick wall and were textured, palisade-like, and villar protrusions had a finger-shaped outline that arose from the cyst wall. These protrusions were interwoven with microtubules in the core and showed small invaginations on their surface; the microtubules did not penetrate into the ground substance. Type 2 sarcocysts had a relatively thin wall and showed minute, naplike elevations on the surface; villar protrusions arose from the cyst walls that were flat and mushroom-like with granules in their core. Apparently, finding sarcocysts in top carnivores is not uncommon, but no other information on these, apparently different, Sarcocystis species has not been forthcoming, to our knowledge, so they must remain species inquirendae. Definitive host: Chrysocyon brachyurus (Illiger, 1815), Maned Wolf. Remarks: Gilioli and Silva (2000) looked at fecal samples of captive maned wolves from 11 zoos in the state of São Paulo, Brazil; they found 6/31 (19%) wolves had oocysts in their feces that they identified only as Cryptosporidium sp. Definitive host: Chrysocyon brachyurus (Illiger, 1815), Maned Wolf. Remarks: Gilioli and Silva (2000) looked in fecal samples of captive maned wolves from 11 zoos in the state of São Paulo, Brazil; they found 3/31 (10%) wolves had oocysts in their feces that they identified only as Eimeria sp. Definitive host: Chrysocyon brachyurus (Illiger, 1815), Maned Wolf. Remarks: Gilioli and Silva (2000) looked in fecal samples of captive maned wolves from 11 zoos in the state of São Paulo, and found that 1/31 (3%) wolves had oocysts in their feces that they determined only as Isospora sp. Definitive host: Chrysocyon brachyurus (Illiger, 1815), Maned Wolf. Intermediate host: Unknown. (2000) looked in fecal samples of captive maned wolves from 11 zoos in the state of São Paulo, and found that 9/31 (29%) wolves had oocysts/sporocysts in their feces that they determined only as Sarcocystis sp. Park and Wildlife Sanctuary in Tamil Nadu and Tadoba National Park, in both areas chital is the most predominant ungulate species. They reported that chital is the intermediate host for Sarcocystis species in dhole. Dhole scats were sampled from 1998 to 2001, and 184/239 (77%) scats from Tadoba and 161/209 (77%) scats from Mudumalai were positive for sporocysts of a Sarcocystis species. Sporocysts measured L × W: 16 × 10, L/W ratio: 1.6, and it was common for densities of 5,000-10,000, up to 26,000, sporocysts/g of feces. Skeletal and heart muscles of chital killed by dhole, or found dead due to other causes, were collected for histopathological examination, but Jog et al. (2003) never definitively stated the number of tissue samples of chital they examined for sarcocysts. They only said that sarcocysts were small, usually <1 mm long, they were not compartmentalized, and "there was no distinct cyst wall" (?). They also said the sarcocysts were found in large numbers in the heart (prevalence 50%) and skeletal muscles (19.5%) collected from Tadoba, whereas in samples collected from Mudumalai, the prevalences were 45% and 48%, respectively. Because neither chital nor dhole are available for experimentation in India, they were unable to demonstrate the life cycle directly; however, Jog et al. (2003) were convinced of the chital-dhole life cycle because of: (1) the consistent occurrence of sarcocysts in chital and sporocysts in dhole; (2) the high proportions of chital among dhole kills; (3) the absence or very low prevalence of sporocysts in other carnivores in the region; and (4) the failure to infect domestic dogs via feeding them chital sarcocysts. Jog et al. (2005) used the same data from their 2003 paper to investigate ecological and coevolutionary aspects of this relationship. Importantly, with no more information than they provided in their 2003 paper, or addressing the guidelines for naming new species, they named the coccidian, Sarcocystis axicuonis. Results of their analyses indicated that sarcocyst density in heart muscles of dhole kills was greater than in chital that died of other causes, and density of sarcocysts in skeletal muscle did not differ between dhole kills and nondhole kills. Jog et al. (2005) further argued that, if Sarcocystis infection in chital does not alter the probability of death due to other causes, then the effect of infection increased the probability of death due to dhole predation. Large numbers of cysts in heart muscle may negatively affect chital stamina making infected chital more susceptible to pack predators (dhole), than stalkers (tigers and leopards), which reinforces the parasite-prey and parasite-host relationships in this system. Jog et al. (2005) argued that if dhole kill more infected than uninfected chital, they are "benefited" by the parasite but, is there a cost to dhole to effectively disseminate the parasite? Dhole scat with large numbers of sporocysts was not diarrheic or otherwise abnormal, which suggested to them that infection in dhole is not pathogenic but no quantitative assessment was conducted. They noted that parasites were overdispersed in the dhole pack (i.e., only a few pack members were passing large numbers of sporocysts at any time). From these observations, they made the interesting suggestion that if carrying the parasite negatively impacts hunting efficiency, then uninfected dhole may do most of the hunting, and if consistent, the "cost" would be negligible to the dhole pack. Maintaining this division of labor between infected and uninfected dhole would be maintained if infected dhole preferentially ate heart tissue with high densities of cysts, and uninfected dhole ate other tissues with no/low densities. Unfortunately, they noted there are no parasite-free dhole-chital populations to serve as a control to estimate actual costs of infection in the two hosts, thus precluding demonstration of cost-benefit analysis of Sarcocystis infection to dhole. If readers are interested, Jog and Watve (2005) further developed their theoretical examination and modeling of parasite-host coevolution and development of mutualistic interactions using the dhole-chital Sarcocystis model. Unfortunately, Jog et al. (2003 , 2005 never completed the basic process of properly naming the Sarcocystis they studied, so it must remain a species inquirenda. Definitive host: Lycaon pictus (Temminck, 1820), African Wild Dog. Intermediate host: Unknown. Remarks: Flacke et al. (2010) surveyed L. pictus in the KwaZulu-Natal province, South Africa, for parasites; they found sporocysts of a Sarcocystis sp. in 12/12 (100%) fecal samples from L. pictus but gave no other information. Knowledge on the parasites of this dog could prove important to its future survival because L. pictus is currently the most endangered carnivore in South Africa, with its total population estimated to be only 300-400 individuals. Definitive host: Lycaon pictus (Temminck, 1820), African Wild Dog. Intermediate host: Unknown. Remarks: Penzhorn et al. (1998) euthanized and necropsied a terminally ill, subadult female L. pictus, and isolated scrapings of the intestinal mucosa; thin-walled sporulated oocysts and many free sporocysts were found, consistent with Sarcocystis species. Two oocysts measured 21 × 15 and 19 × 16. Sporocysts were L × W (n = 12): 15.7 × 10.2 (14-17 × 9-11), L/W ratio: 1.5; and SZ (n = 4) measured 9-11 × 2 (10.3 × 2). On histological examination, they saw large numbers of thin-walled, sporulated oocysts, each with two sporocysts and four crescent-shaped SZ with prominent N in their caudal third. Oocysts were present in the lamina propria at the tips of almost all small intestinal villi but there was no evidence of inflammatory reaction or tissue necrosis. Definitive host: Unknown. Intermediate host: Nyctereutes procyonoides (Gray, 1834) , Raccoon Dog. Remarks: Levine and Ivens (1981) listed this form and said that sarcocysts were found in the muscles of the raccoon dog in Primorye (former USSR). However, neither Pellérdy (1974) nor Levine and Ivens (1981) , nor Levine (1986) Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Willingham et al. (1996) surveyed the feces of road-killed red foxes in metropolitan Copenhagen, Denmark; "coccidia oocysts" were detected in 2/68 (3%) foxes, mostly juveniles, but no other information was given. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Sturdee et al. (1999) detected what they said was C. parvum in 22/184 (12%) red fox fecal samples tested with a genus-specific monoclonal antibody for C. parvum. Their results purportedly emphasized the widespread distribution of Cryptosporidium among wild mammals in Britain and allowed them to suggest the potential for transmission between wild mammals, via direct exposure, to those using the countryside for professional or recreational purposes (e.g., farmers and ramblers). Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Zhou et al. (2004) collected the feces of 471 wild mammals from four counties in the Chesapeake Bay area of Maryland, USA, and found 6/76 foxes (8%), presumably V. vulpes (?) (the host binomial was not stated), to be infected with the C. canis fox genotype (4), the C. canis dog genotype (1), and a Cryptosporidium muskrat genotype (1). The species and genotypic nature of Cryptosporidium in each fecal sample was determined by a PCR-restriction fragment length polymorphism (RFLP) method based on the small-subunit rRNA gene. Original host: Vulpes (syn. Alopex) lagopus (L., 1758), Arctic or Blue Fox. Remarks: Nukerbaeva and Svanbaev (1973) said they found this form in 30/1,089 (3%) Arctic foxes in Kazakhstan of the former USSR. Their oocysts were ellipsoidal, 14 × 10 (13-15 × 8-11), L/W ratio: 1.4, with a 2-layered wall, ∼1 thick; OR: present; M, PG: both absent. Sporocysts were ellipsoidal, 6-7 × 3-4, SB, SSB, PSB, SR: all absent. Nukerbaeva and Svanbaev (1973) presented a modest line drawing, but little structural data and this species has not been seen since its original description. Original host: Vulpes (syn. Alopex) lagopus (L., 1758), Arctic or Blue Fox. Remarks: Rastégaïeff (1929c) published a note préliminaire naming this "species" from oocysts in the feces of an Arctic fox collected in Murmansk Oblast, northern Russia, on the south side of the Barents Sea. Oocysts were spheroidal to ovoidal, 18 × 11-14, with a 1-layered wall and a M occupying the entire small end of the oocyst, which lacked an OR. Sporocysts were ellipsoidal. Pellérdy (1974a) questioned whether this was a real species (?) of the fox, and Levine and Ivens (1981) thought it looked like a rabbit coccidium. Rastégaïeff (1930) said she found it in the same fox on both October 10 and November 19, 1928. Levine and Ivens (1981) indicated they had seen a very poor line drawing, but we have a copy of her (1929) preliminary note, which does not have a line drawing. Unfortunately, she named the species in honor of Professor F. Mesnil, Laboratory of the Parasitology School, Leningrad, but this can only be considered a species inquirenda. Original host: Vulpes vulpes (L., 1758), (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Golemansky and Ridzhakov (1975) said they found oocysts of an unnamed Eimeria sp. in 6/146 (4%) red foxes in Bulgaria with oocysts that were L × W: 33 × 20 (28-38 × 17-23) and had sporocysts that measured 13-15 × 8-10, without a SB. They said these likely were a pseudoparasite of the fox, probably oocysts of a rabbit species just passing through the gut of the fox. We agree. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Magi et al. (2009) examined fecal samples of red foxes in Tuscany (Cecina, Grosseto, Pisa, Siena), central Italy. Samples were examined by "coprological methods and microscopy," and "oocysts of Eimeria spp." were detected in 10/110 (9%) samples. Original host: Vulpes vulpes (L., 1758), (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Galli-Valerio (1929a) found this form in V. vulgaris collected at 1,650 m, in the Val de Bagnes, Switzerland; oocysts were ovoidal, with one end "barely flattened" as a nearly invisible M, measured 17 × 14, with ovoidal sporocysts, each 6 × 4.5, and contained two SZ, ∼4 × 2.4. A year later he found this species in another V. vulgaris collected on Fignards Mountain near Torgon, Switzerland (Galli-Valerio, 1930) . Watkins and Harvey (1942) reported this species in 11/52 (21%) silver fox cubs dying in England and reportedly found it in ∼10% of the adults and perhaps 25% of the fox cubs from 15 fox farms in England. Svanbaev (1960) reported it in 4/18 (22%) silver foxes in the Alma Atinsk Oblast, Kazakhstan and said the oocysts sporulated in 3-4 days at 25°C in 2% K 2 Cr 2 O 7 solution. Golemansky and Ridzhakov (1975) said they found oocysts of this species in the feces of 15/146 (10%) foxes in Bulgaria. Combining descriptive features from these other authors, the sporulated oocysts are ovoidal; number of walls, 1 (?); wall characteristics: smooth, colorless, 0.8-1.5 thick; L × W: 17 × 14 (16-26 × 12-24), L/W ratio: 1.2; M; barely visible or absent; OR, PG: both absent. Likewise, sporocysts may be ovoidal to ellipsoidal; L × W: 6 × 4.5 (5-6 × 3-6); L/W ratio: 1.3; SB, SSB, PSB: all absent; SR: present; SZ: comma-shaped, 4-5 × 2. Frank (1978) said she found this species in V. vulpes trapped near Lake Neusiedl, Austria, on the border with Hungary. Nonetheless, no published description to date has provided either a line drawing or a photomicrograph of a sporulated oocyst. Thus, this form must be relegated as a species inquirenda. Original hosts: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox; Vulpes (syn. Alopex) lagopus (L., 1758), Arctic or Blue Fox. Remarks: Dahlgren and Gjerde (2010b) infected six of each fox species with fresh portions of esophagus, diaphragm, and abdominal muscles from moose, Alces alces, from Norway. They reported unsporulated, subspheroidal Hammondia oocysts were found in the mucus along the entire posterior half of the small intestine in one of each species of fox killed on the 7th DPI and a few Hammondia oocysts also found in the ileum of one silver fox killed on 14 DPI. Oocysts were shed in large numbers during the first 1-5 days of patency, then in small numbers or intermittently thereafter; patency began ∼13-14 DPI after foxes ingested moose meat. Hammondia oocysts were detected microscopically in 11/12 foxes that ingested moose flesh (their Table 2 , p. 1552), and samples from 7 foxes, 4 V. vulpes and 3 V. lagopus, were positive on agarose gels after PCR using Hammondia-specific primers. Moose previously were reported to be an intermediate host of H. heydorni infecting dogs (Dubey and Williams, 1980) , but this was the first report of moose as intermediate host of a Hammondia species infecting foxes. Citing recent molecular comparisons of Hammondia isolates from dogs and foxes, Dahlgren and Gjerde (2010b) suggested that the Hammondia oocysts they found in red and arctic foxes in Norway might be different from H. heydorni known from dogs. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Dahlgren and Gjerde (2010b) found oocysts in the feces of one fox, on the 8th DPI, which measured ∼35 × 25, and "resembled those of Isospora canivelocis." No other information was given. Original host: Vulpes vulpes (L., 1758), (syn. V. vulgaris Oken, 1816), Red or Silver Fox. Remarks: Stuart et al. (2013) surveyed fecal samples of red foxes killed throughout Ireland, and "Isospora-like oocysts" were seen in 8/91 (9%) samples, but no other information was given. Synonym: Eimeria vulpes Patnaik and Acharjyo, lapsus. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Remarks: Galli-Valerio (1931) measured oocysts that were 25 × 24, with a visible M, and two sporocysts each with four SZ, but gave no other measurements or qualitative descriptive information and then named this "species" as new! His description was less than marginal and he did not provide a line drawing or photomicrograph as a type. Nonetheless, Svanbaev and Rachmatullina (1971) said they found it in 6/85 (7%) common foxes (V. vulpes). The oocysts they studied were ovoidal (25-28 × 20-22) or spheroidal (25-28 wide), light gray, with a bilayered smooth wall, ∼2 thick, and M, OR, and PG all absent. Sporocysts were ovoidal (14-17 × 11-14), without SB, SSB, and PSB. Golemansky and Ridzhakov (1975) measured oocysts that were 20-23 × 18-21. The measurements by Galli-Valerio (1931) suggest that many of the oocysts measured were end-on views resulting in the mean of the measurements to be erroneously more spheroidal. It is possible that this is Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi, but the oocyst sizes are somewhat larger suggesting Isospora vulpis Galli-Valerio, 1931 may represent a separate species. The exception is Golemansky and Ridzhakov (1975) , who may indeed have seen a member of the Isospora ohioensis-complex. It is likely that Bledsoe (1976a,b) actually was working with I. (=C.) vulpis rather than I. (=C.) vulpina. However, Galli-Valerio (1931) stated that the oocysts had a micropyle and his measurements suggest subspheroidal rather than ellipsoidal oocysts. It is likely that he was in error on both accounts and that the description by Svanbaev and Rachmatullina (1971) is the most accurate. Unfortunately, however, no published description to date has provided either a line drawing or a photomicrograph of a sporulated oocyst. Thus, this form must be relegated to a species inquirenda. Synonym: Hammondia sp. Ashford, 1977. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Remarks: Levine and Ivens (1981) placed this form in the genus Isospora, but it is probably best left a species inquirenda. Ashford (1977) found some oocysts, 14 × 12 in the feces of 1/22 (4.5%) red foxes in England. He (Ashford, 1977) said the oocysts resembled those of "Hammondia" hammondi, but they were not infective for mice so he suggested they might belong to another species of Hammondia. Levine and Ivens (1981) thought it could just as well be an Isospora or a Besnoitia species. It is probably best relegated to a species inquirenda. Original host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Remarks: Golemansky and Ridzhakov (1975) found oocysts in 2/146 (1%) red foxes in Bulgaria and measured a few; oocysts were L × W: 38 × 33 (30-52 × 25-35) and contained 5-16 spheroidal sporocysts that were about 11-12 wide, without a SB, but with a SR consisting of many residual granules. They likely had found oocysts of an Adelina sp. from an arthropod or annelid and this was a spurious finding. Definitive host: Unknown. Intermediate host: Vulpes corsac (L., 1758), Corsac Fox. Remarks: Levine (1986) listed this species as a valid, named Sarcocystis species, but gave no other information. Odening (1998) said that Pak (1979) named this species from sarcocysts found in the corsac fox, but specimens were not preserved so follow-up molecular examination of the specimens cannot be done. Gjerde and Schulze (2014) suggested that because the specimen described is unrecognizable today, it should be considered a species inquirenda and its name should become a nomina dubia. Neither Levine (1986) nor the three of us were able to secure a copy of the original paper, so we must concur with the opinion of Gjerde and Schulze (2014). We do, however, list its complete citation in our References. 's (1989b) classification of 24 sarcocyst-types to identify a few sarcocysts of this form (type 10 villar protrusions) in one semi-thin tissue section of diaphragm from a sheep in Sicily, Italy. Sarcocysts were 800 × 300 and divided into compartments by septa. The cyst wall was ∼5 thick, with radial striations. Villar protrusions had microtubules that extended from the apex to the base of the villi, all characteristics of S. gracilis from roe deer. However, this identification is only circumstantial, and this form should be regarded as a species inquirenda. We mention this form here because we know from recent work (Gjerde, 2012) that S. gracilis sarcocysts in roe deer muscle can be transmitted to both V. vulpes and V. lagopus, which then shed sporocysts in their feces 9 DPI with infected deer flesh. Definitive hosts: Vulpes (syn. Alopex) lagopus (L., 1758), Arctic or Blue Fox; Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Intermediate host: Bos taurus L., 1758, Aurochs. Remarks: Odening et al. (1996a) found sarcocysts in the musculature of a dwarf zebu born in a German zoo. One of the four forms they found, "mostly resembles Sarcocystis gracilis Rátz, 1909 from roe deer." We mention this form here because we know from recent work (Gjerde, 2012) that S. gracilis sarcocysts in roe deer muscle can be transmitted to both V. vulpes and V. lagopus, which then shed sporocysts in their feces 9 DPI with infected deer flesh. Obviously, this form must remain a species inquirenda because no other data were given to confirm its identification as S. gracilis in the dwarf zebu. Remarks: Odening (1998) mentioned this species as valid in his compilation of 189 Sarcocystis names and said it was similar to S. citellivulpes. However, the authors are cited only in a Russian reference book (Anon, 1984) that is unavailable to us. Odening (1998) gave such little detail in his citation, that we think it is best, at this time, to relegate this form to a species inquirenda. To our knowledge, there have been no other references to this species since 1984, other than by Odening (1998). Definitive host: Unknown. Intermediate host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Remarks: Odening (1998) , who listed this as a valid Sarcocystis species name, said that Pak et al. (1991) named it from sarcocysts found in the red fox, but specimens were not preserved so followup molecular examination of specimens cannot be done. Dubey et al. (2015a) listed the form by Pak (1991) from the red fox as Sarcocystis sp. Gjerde and Schulze (2014) suggested that because the specimen described is unrecognizable today, it should be considered a species inquirenda and its name should be a nomina dubia. We were not able to secure a copy of the original paper, so we must concur with the opinion of Gjerde and Schulze (2014). Intermediate host: Capra ibex L., 1758, Alpine Ibex. Remarks: Biocca et al. (1975) found sporocysts in the feces of "some" of 12 red foxes in the Gran Parasiso National Park (GPNP) in Italy; these sporocysts measured 13-15 × 8-10, with a large SR, but no SB. They also found an ibex with sarcocysts in its muscles. Levine and Ivens (1981) reported that Biocca et al. (1975) infected both the timber wolf and domestic dog by feeding them sarcocyst-infected esophageal, heart, diaphragm, intercostal, and abdominal muscles from the infected ibex. The prepatent period in the fox was 11 days, in the wolf it was 12 days, and in the domestic dog it was 20 days; the patent period in these hosts was 62, 67, and 66 days, respectively. They also tried, unsuccessfully, to infect a domestic cat, a lion, a ferret, and a kestrel (Falco tinnunculus) with sarcocysts from the ibex, but none of them passed sporocysts. However, there is no evidence or correlation that the sporocysts in the foxes were the same species as the sarcocysts in the ibex. Later, Cornaglia et al. (1998) described the ultrastructural morphology of the sarcocyst wall in the muscles from an ibex found dead in the same national park in Italy. They also took samples of the diaphragm from 52 ibexes of different age and sex, from the GPNP. The morphology of the cyst wall led to the identification of three types of sarcocysts in the ibex of the GPNP and a further type in one Hispanic ibex was also collected. Cornaglia et al. (1998) concluded, "the morphology of the sarcocysts' walls was similar to the wall of the species described in the domestic ruminants from several authors." Only one Japanese red fox had sarcocysts in its muscles and no inflammatory host response was associated with the sarcocysts. Ultrastructurally, the sarcocyst wall was thin and showed minute undulations. Kubo et al. (2009) said that these sarcocysts were similar to sarcocysts they found in the Japanese raccoon dog (N. procyonoides viverrinus) and Japanese martens (M. m. melampus) during the same survey. This is the first published report of muscular sarcocystosis in Japanese carnivores. There was no attempt to identify this species so this form must remain a species inquirenda. Definitive host: Vulpes vulpes (L., 1758) (syn. V. vulgaris Oken, 1816), Silver or Red Fox. Intermediate host: Unknown. Remarks: Golemansky (1975b) found free oocysts and sporocysts in the feces of 14/146 (10%) V. vulpes in Soria, Bulgaria. Oocysts measured L × W: 18.6 × 14.0 (17-20 × 10-18) and sporocysts were 15.3 × 10.2 (13-18 × 9.5-11). He thought these oocysts/sporocysts resembled those of S. fusiformis and S. tenella, but the only conclusion he drew was "the European common red fox is one of the vectors in the maintenance and distribution of sarcosporidiosis in nature." This can only be considered species inquirenda. 2) using TEM and saw it had thin-walled sarcocysts that were 0.5-0.7 thick, lacking protrusions, and exhibiting minute type 1 (Dubey et al., 1989b) undulations on the wall. Two of these sarcocysts measured 95 × 60 and 110 × 65. Under TEM the sarcocyst wall had minute, electron-dense undulations located at irregular intervals. Only bradyzoites were seen and three of them, in longitudinal section, were 5.0-5.7 × 1.6-1.9. Rhoptries were prominent, their bulbous blind end was sometimes turned toward the conoidal end, and their micronemes were in the anterior half of the bradyzoite; all these features making this species ultrastructurally distinct from S. neurona. However, to our knowledge it has never formally been named. Original host: Lontra canadensis (Schreber, 1777), North American River Otter. Remarks: Gaydos et al. (2007b) collected feces from L. canadensis living along the Puget Sound Georgia Basin (PSGB) marine ecosystem between Washington state, USA, and the southern tip of Vancouver Island, British Columbia, Canada, to look for the presence of Cryptosporidium and Giardia oocysts and cysts, respectively. In Washington state, they collected fecal samples from 13 locations in PSGB, and off Vancouver Island, from 30 locations; they found 4/57 (7%) samples from 4 locations in Washington state, and 5/36 (14%) fecal samples from 2 locations on Vancouver to have Cryptosporidium oocysts in their fecal material. Overall, they found Cryptosporidium oocysts in 9/93 (10%) fecal samples from river otters in the PSGB. Even when some parasites, such as Cryptosporidium, are only identified to genus in wildlife surveys, we can see that there are so many examples of the potential for these and other parasites to be transmitted between wildlife and humans. Definitive host: Unknown. Intermediate host: Lutra lutra (L., 1758), European Otter. Remarks: Wahlström et al. (1999) found sarcocysts in the skeletal muscle of one L. lutra raised in Norway but had died in captivity in Sweden. The sarcocysts were 0.3-2.3 mm long and 0.05-0.25 mm wide. Under LM, sarcocyst walls were thin, <3 μm, and had a serrated surface, but did not have visible projections. By TEM, the sarcocyst wall was 0.6-1.8 thick and had minute undulations covering its entire surface, giving the wall a wavy appearance. Septa were indistinct and the sarcocysts had few metrocytes, but many bradyzoites. These sarcocysts were found in only 1/70 (1%) otters subjected to necropsy in Sweden. Original host: Martes zibellina (L., 1758), Sable. Remarks: Matschoulsky (1947a,b) reported three species of coccidia in 32/144 (22%) sables, "two already known and one is new." In one sable he found oocysts that were round/ovoidal and surrounded by a 2-layered wall, ∼1 thick, without a M. Ovoidal oocysts were L × W: 13.1 × 11.3 (12-14 × 10-12), L/W ratio, 1.2. Although he could not get oocysts of this "species" to sporulate, he said the defining feature of this species is its small size, which "differs drastically from coccidia that were found in sables before. That is why we name this species as new." Fortunately, he did not give it a name, and because he never saw sporulated oocysts, it cannot be placed into a genus. Original host: Martes foina (Erxleben, 1777), Beech Marten. Remarks: Rademacher et al. (1999) reported temporary episodes of diarrhea in four captive beech martens that were not related to each other, and they found numerous oocysts of a Cryptosporidium sp. in the feces of all of them. The oocysts measured, L × W (n = 30): 3.6 × 3.1 (3-4 × 3-4); L/W ratio: 1.2, but no other information was given. Original host: Martes martes (L., 1758), European Pine Marten. Remarks: Yakimoff and Gousseff (1934) Original host: Martes melampus (Wagner, 1840), Japanese Marten. Remarks: Yanai et al. (1995) studied Japanese martens in Gifu prefecture, Japan and found nodular lesions containing meronts and merozoite-gametocytes of a Hepatozoon species in 67/70 (96%) wild martens. The heart was the most commonly parasitized organ (67/70, 96%) followed by perirenal adipose tissue (25/70, 36%), diaphragm (9/58, 16%), mesentery (10/68, 14%), tongue (1/7, 14%), omentum (8/57, 14%), and perisplenic adipose tissues (7/70, 10%). Two types of nodular lesions were seen, each based on different developmental stages: nodules containing meronts, and nodules that consisted of an accumulation of phagocytes containing merozoites or gamonts. Nodules containing meronts were 50-400 wide. Yanai et al. (1995) studied mature meronts and membrane-bound merozoites with LM and TEM but chose not to name the parasite. Definitive host: Martes pennanti (Erxleben, 1777), Fisher. Intermediate host: Unknown. Remarks: Dubey (1982b) surveyed feces for coccid ians from a variety of wild carnivores in Montana, USA and reported sporocysts of Sarcocystis in the feces of 1/6 (17%) fishers. Sporocysts from this host varied from 12.5-13.0 × 8.5-9.0 (n = 3). No attempt was made to identify the species beyond genus; thus, a species inquirenda. Definitive host: Unknown. Intermediate host: Martes pennanti (Erxleben, 1777), Fisher. Remarks: Gerhold et al. (2005 described meningoencephalitis due to S. neurona in a fisher from Maryland, USA; they also found intramuscular sarcocysts "of a possibly unrecognized Sarcocystis species." Although the structure and ultrastructure of the muscle sarcocysts from the fisher were similar to those of S. neurona, they were unable to amplify S. neurona DNA from these muscle forms; they said this may be related to technical difficulties, or this organism may be an unrecognized species that has sarcocysts morphologically similar to those of S. neurona. Definitive host: Unknown. Intermediate host: Martes melampus melampus Gray, 1865, Japanese Marten. Remarks: Kubo et al. (2009) examined 65 wild carnivores on Honshu, Japan for sarcocysts and found three Japanese red martens had sarcocysts in their muscles, and no inflammatory host response was associated with the sarcocysts. Ultrastructurally, the sarcocyst wall was thin and showed minute undulations. Kubo et al. (2009) said these sarcocysts were similar to sarcocysts they found in the Japanese raccoon dog (N. procyonoides viverrinus) and the Japanese red fox (V. v. japonica) during the same survey. This was the first published report of muscular sarcocystosis in Japanese carnivores. There was no attempt to identify the species; thus, a species inquirenda. Original host: Meles anakuma Temminck, 1844, Japanese Badger. Remarks: Kamiya and Suzuki (1975) examined the preserved intestine of one badger, mostly for trematodes, but when they looked at tissue sections of the jejunum mucosa they saw endogenous stages of a coccidium represented by macro-and microgameto cytes and oocysts. Microgametocytes with many microgametes were ∼20 wide, macrogametocytes with many basophilic granules were 15-22 × 13-20, and unsporulated oocysts were 20-21 × 14-17. Although they concluded these stages represented an Eimeria species there is no way to know that from the few tissue stages measured and unsporulated oocysts. Definitive host: Unknown. Intermediate host: Meles anakuma Temminck, 1844, Japanese Badger. Remarks: Kubo et al. (2009) examined 65 freeliving carnivores on Honshu, Japan for muscular Sarcocystis and two Japanese badgers had sarcocysts in their muscles, but no inflammatory host response was associated with them. Ultrastructurally, the sarcocyst wall was thick with numerous finger-like protrusions that contained microtubules. Kubo et al. (2009) said these sarcocysts were not similar to sarcocysts they found in the Japanese raccoon dog (N. p. viverrinus), the Japanese red fox (V. v. japonica), and the Japanese marten (M. m. melampus) during the same survey. This was the first published report of muscular sarcocystosis in Japanese carnivores, but there was no attempt to identify it to species, so this form must remain a species inquirenda. Definitive host: Unknown. Intermediate host: Meles meles (L., 1758), European Badger. Remarks: Odening et al. (1994a) found a dead female M. meles on the road, ∼50 km northeast of Berlin, Germany. Macroscopic sarcocysts were found in the tongue and, after fixation and sectioning, they were examined through TEM. Sarcocysts were up to 1.4 mm long, with a maximum width of 185 μm, and had compartments of various sizes, with bradyzoites that measured L × W (n = 30): 12.3 × 3.1 (11-13 × 2.8-3.5). Odening et al. (1994a) said, "No sarcocysts from mustelids have been described by electron microscopy so far," but they obviously missed the paper by Cawthorn et al. (1983) , who used TEM to look at sarcocysts in experimentallyinfected Richardson's ground squirrels after they had fed on Sarcocystis sporocysts recovered from the American badger, Taxidea taxus (Schreber, 1777). Odening et al. (1994a) compared their form to similar sarcocysts studied from roe deer by Entzeroth (1982a,b), and Sugár et al. (1990) , and said, "the similarity of the…forms from roe deer with each other and with the form from the European badger is so great, that we can regard all these forms as most likely being identical." Without more substantive data, this form must remain a species inquirenda. Synonym: cf. Sarcocystis sebeki of Tadros and Laarman, 1976. Definitive host: Unknown. Intermediate host: Meles meles (L., 1758), European Badger. Remarks: Odening et al. (1994b) found four dead M. meles (Nos. 3, 2, 6, 9; 1 male and 3 females, respectively) on various roads, 45-75 km from Berlin, Germany. Sarcocysts were found in the tongue, thigh, loin, and thorax and, after fixation and sectioning, they were examined via LM and TEM. Sarcocysts were 6.5-9.0 mm long, and 172-200 μm wide in the fresh state, and their bradyzoites were squat and fusiform, 6.9 × 1.9 (6.1-7.2 × 1.6-2.2). In semi-thin sections bradyzoites were 5.9 × 2.0 (5.7-6.3 × 1.7-2.2). TEM showed a cyst wall 0.9-1.4 thick with no protrusions. Small elevations of the primary cyst wall were 0.08-0.09 long, the fossule-like invaginations in-between had a maximum diameter of 0.05. These small elevations and invaginations were underlayed with an osmiophilic layer. The authors (1994b) "assigned" this form to be near S. sebeki (cf. Tadros and Laarman, 1976 , 1978 , 1979 , 1980a ,b, 1982 "because it is morphologically very similar" to it. Definitive host: Unknown. Intermediate host: Millivora capensis (Schreber, 1776), Honey Badger. Remarks: According to Levine and Ivens (1981) , Viljoen (1921) found sarcocysts in the striated muscles of a honey badger, but they did not see his actual paper and got the citation from Nietz (1965). Levine (1986) did not list this species, and Dubey et al. (2015a) list neither the Viljoen nor Nietz references and make no mention of a Sarcocystis species in the honey badger. Viljoen (1921) does not retrieve in Google Scholar and we were unable to obtain a copy of the Nietz paper. Remarks: Blankenship-Paris et al. (1993) examined a 4-month-old ferret that was ill, dehydrated, and had pasty, dark feces. The ferret was housed with another sibling and its dam, but neither showed similar signs, and repeated fecal exams of both were negative for coccidial oocysts. Also, fecal exams of other members of the colony and necropsy of eight other ferrets in the colony did not demonstrate any coccidia. However, when the 4-month-old ferret was necropsied, Blankenship-Paris et al. (1993) reported that "enteric coccidiosis" was the cause of its illness, based on histological findings of some endogenous stages. No other identification was made. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks: Bronson et al. (2007) examined mortality, retrospectively, based on the pathology records of 107 captive animals held at the Smithsonian's National Zoological Park, Washington, D.C., USA from 1989 to 2004. They said that "the most common cause of death among juvenile ferrets was gastrointestinal disease, found in 11/21 (52%)," with seven of those cases caused by coccidiosis. No identity of the coccidian nor other information was given. Original host: Neovison vison (Schreber, 1777) (syn. Mustela vison L., 1766), American Mink. Remarks: Davis et al. (1953) were among the first to notice liver coccidiosis in a mink, and they described characteristic pathological and anatomical changes caused by this parasite, but because they examined only visceral organs, they did not identify the coccidian. Original host: Mustela nigripes (Audubon and Bachman, 1851), Black-footed Ferret. Remarks: Jolley et al. (1994) reported seeing meronts and oocysts of a small coccidian in the cells lining the trachea, a bronchus, and in associated bronchial glands in one black-footed ferret, which also had canine distemper. Because no sporulated oocysts could be found they could not name or characterize it further. No lesions in tracheal tissue could be attributed to the presence of this parasite. Original host: Mustela nigripes (Audubon and Bachman, 1851), Black-footed Ferret. Remarks: Jolley et al. (1994) reported seeing merozoites of another unidentified coccidian in an impression smear of the epithelium of the urinary bladder of the same ferret in which they saw the respiratory coccidian stages (above). No lesions in bladder tissue could be attributed to the presence of this parasite, but they cautioned that this form would need differentiation from Toxoplasma, Neospora, and Hepatozoan species, which it resembles in size, ability to invade a variety of tissue types, and merogonic development. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks: Li et al., 1996 described proliferative bowel disease in 4/19 (21%) ferrets they found to be coinfected with a Desulfovibrio sp. (gram negative, comma-to spiral-shaped bacteria, often associated with proliferative bowel disease) and coccidia. Thick, rigid colons, and enlarged mesenteric lymph nodes were palpable in all four, which had a history of lethargy, anorexia, weight loss, and diarrhea. These ferrets were dehydrated and emaciated, and one had occult blood in its feces whereas another had a prolapsed rectum. Tissue sections showed proliferative changes that included a marked increase in mucosal thickness, and glandular or crypt length and irregularity with pseudovillus formation. Epithelial cells were markedly hyperplastic, hyperchromatic, and piled on each other with numerous mitotic figures. Inflammatory cells consisted predominantly of neutrophils (two ferrets) or lymphocytes and macrophages (two ferrets), and some eosinophils in all four. They described the coccidial organisms to consist predominantly of meronts and merozoite stages. Many coccidial stages were seen within the apical cytoplasm of hyperplastic epithelial cells, but feces were not examined and no attempt was made to identify the organism other than to call them coccidia. They mentioned that the mechanism by which Desulfovibrio sp. and its relatives induce proliferative bowel disease is unknown, but "the presence of a second organism may in some instances be required for ICOs to cause intestinal lesions." Original host: Mustela (syn. Putorius) putorius furo L., 1758, Domestic Ferret. Remarks: Gómez-Villamandos et al. (1995) reported that four of five ferrets shipped to their department in Córdoba, Spain became ill several days after arrival showing anorexia, depression, and yellow diarrhea; they died 48-72 hours after onset of the illness began. The ferrets were nonpregnant females weighing 500-550 g and originated on a goat farm. Tissue samples were fixed and prepared for LM and TEM. Microscopic examinations of the intestinal tract of all ferrets showed numerous spheroidal to ovoidal bodies, in various stages of development, attached to the brush border of enterocytes and cryptal epithelial cells of the distal large intestine. Villous atrophy was severe and a cellular infiltrate composed of neutrophils, eosinophils, and macrophages was seen in the submucosa, whereas desquamated epithelial cells with parasitic stages were seen in the intestinal lumen. All stages of the life cycle were observed including trophozoites, two types of meronts with their merozoites, macro-and microgametes, and unsporulated and sporulated oocysts. No attempt was made to name the parasite. Original host: Mustela (syn. Putorius) putorius furo L., 1758, Domestic Ferret. Remarks: Rehg et al. (1988) identified the first case of Cryptosporidium in ferrets. Two unrelated young ferrets, 4-and 8-months-old, died from unknown causes on the same day. They made stained fecal smears, and then took sections of both gastrointestinal (GI) tracts and stained paraffin-embedded sections with hematoxylin-eosin for the histopathology; they also fixed, cut, and stained the GI with uranyl acetate and lead citrate for TEM. Fecal smears showed Cryptosporidium oocysts that were ∼3-5. Histological sections demonstrated cryptosporidial stages, 2-5 wide, associated with the brush border of the epithelial cells at the tips and lateral margins, but not in the crypts, only in the small intestine (predominantly ileum); no such stages were seen in the gall bladder, trachea, or lungs. Trophozoites, meronts, macrogametes, and oocysts were detected by TEM in the small intestine. Subsequently, examination of fresh fecal samples of the ferret population at the research facility, and of the new arrivals, revealed the presence of Cryptosporidium as a subclinical disease in a high percentage of ferrets. The infection persisted for several weeks both in immunocompetent and immunocompromised (treated with dexamethasone) animals. Fecal smears showed Cryptosporidium oocysts in 9/22 (41%) ferrets at the facility, and in 31/44 (70%) new arrivals. Neither species nor genotypes of the isolates were identified because only LM, TEM, and histopathological methods were used. Original host: Neovison (syn. Mustela) vison (Schreber, 1777), American Mink. Remarks: Very small oocysts, 4-5 long, identified as Cryptosporidium sp. were found in the feces of 11/40 (27.5%) mink pups on 5/13 (38%) fur farms in Iceland, but no further account of this species was ever published (SkírNisson and Pálmadóttir, 1993). Original host: Mustela erminea L., 1758, Ermine. Original host: Mustela erminea L., 1758, Ermine or Stoat. Remarks: The paper of Nukerbaeva and Svanbaev (1972) is unavailable to us or it may not even exist. In their survey of fur animals in Kazakhstan, Nukerbaeva and Svanbaev (1977) said they found 1/7 (14%) stoats infected with E. baskanica Nukerbaeva and Svanbaev, 1972; however, Global Names Index (gni.globalnames. org/) listed the authority as Nukerbaeva and Svanbaev, 1973. In their 1977 paper they briefly mentioned the oocysts of this form to be ovoidal with tapered poles, dimensions of 11.2-12.6 × 8.4-9.8, lacking both M and PG, but having an OR; the sporocysts were described only as ovoidal, with two bean-shaped SZ and an SR, but no mention was made about presence/absence of a SB. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks: Yakimoff and Gousseff (1936) reported finding oocysts in two polecats taken from the Polotsk District, in the Republic of Belarus on the Dvina River. They found only small numbers of perfectly round oocysts with a smooth, double-layered wall, that measured 21.6-23.4 wide; these oocysts lacked M, OR, and PG. The elongate-ovoidal sporocysts were 12.6 × 5.2, and apparently lacked SB, SSB, PSB. They said the oocysts they observed were somewhat similar to those of E. furonis, which they are not (21.6-23.4 vs.12.8 × 12.0). Ultimately, they identified their oocysts as E. mephitidis, a species described from the striped skunk (M. mephitis) from Ohio, USA, by Andrews (1928); the oocysts are similar in size, but those of E. mephitidis have a circular M, which was not seen in those described by Yakimoff and Gousseff (1936) . The line drawing provided by Yakimoff and Gousseff (1936) is strikingly similar to the line drawing of E. hiepei, which was described from the European mink by Gräfner et al. (1967) , but E. hiepei has smaller oocysts (13.5-16.6 wide). Original host: Mustela nigripes (Audubon and Bachman, 1851), Black-footed Ferret. Remarks: Jolley et al. (1994) reported finding oocysts representing three Eimeria species in 6/6 (100%) M. nigripes in Wyoming, USA. Two of the species were identified as E. furonis and E. ictidea. The third form, which they said was seen only rarely, had oocysts that were larger than those of the first two, but they made no attempt to identify or characterize it in any way other than to say it was oval and five oocysts they measured were L × W: 37.0 × 22.3 (35-39 × 21-23), L/W: 1.7. Confirming that this was likely an eimerian they said, "sporulated oocysts of the large oval form were rarely seen," and "attempts to sporulate this species were variably successful under the conditions described, often with little or no development seen after a 10-day incubation period in the potassium dichromate solution. Oocysts taken from the intestinal lumen failed to sporulate, whereas some of those passed in the feces successfully did so." They also pointed out that neither Levine and Ivens (1981) , from mustelids and other carnivores, nor Levine and Ivens (1990) and Thomas and Stanton (1994) from prairie dogs, rock squirrels, or ground squirrels had reported Eimerian oocysts with these measurements. Original host: Mustela nivalis L., 1766, Least Weasel. Remarks: Musaev and Veisov (1983) found oocysts of an Eimeria sp. in 1/43 (2%) weasels they examined; they thought these oocysts were slightly different from those of E. mustelae, but they declined to name their form even though they provided a line drawing. The oocyst wall was smooth, colorless, 1.5-2.0 thick. Sporulated oocysts were L × W (n = 49): 20.3 × 14.8 (17-22 × 12-17), L/W ratio 1.4 (1.2-1.6); M, OR, PG: all absent. Sporocysts were described as ovoidal to pear-shaped although this is not evident in their line drawing. Sporocysts were L × W: 8.5 × 6.5 (6-10 × 4-8); SB, SSB, PSB: all absent (line drawing); SR: present, as a group of fine granules. Sporozoites measured were 5-9 × 3-7 and each had a RB at their broader end. Sporulation time was 48-72 hours in 2.5% potassium dichromate (K 2 Cr 2 O 7 ) solution at 25-30°C. They said the oocysts measured were obtained from the contents of the large intestine and the host was collected in the Batabag Shahbusk region of the Nakhitschevansk, Azerbaijan. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks: Patterson and Fox (2007) reported oocysts they identified as I. ohioensis in fecal samples of healthy domestic ferret kits in a large ferret breeding operation on the same premise as juvenile domestic dogs; they neglected to mention how they arrived at this identification. It is likely their identification was either a spurious finding of a dog oocyst passing through the gut of the ferrets or is a new/different species morphologically indistinguishable from the oocysts of I. (=C.) ohioensis. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks: Bell (1994) reported that Isospora sp. oocysts are commonly shed by ferrets 6-16 weeks of age and is the same species that commonly affects puppies and kittens, but the "species" was not identified. Original host: Mustela (syn. Putorius) putorius furo (L., 1758), Domestic Ferret. Remarks Definitive host: Unknown. Intermediate host: Mustela nivalis L., 1766, Least Weasel. Remarks: Tadros and Laarman (1979) reported muscular sarcosporidiosis for the first time from the common European weasel in the Netherlands. They examined the morphology of sarcocysts both in fresh and stained histological preparations. The sarcocysts were several mm long by 0.15 mm wide and had a smooth wall without cytophaneres; the cysts were compartmentalized and had metrocytes 3.5 wide and bradyzoites about 9 × 2.5. Levine and Ivens (1981) speculated that these were very similar to the sarcocysts of S. sebeki found in Apodemus sylvaticus (L., 1758). In addition to its muscle sarcocysts, the weasel also had sporocysts of S. putorii in its feces. They attempted to complete the sexual cycle by feeding muscle sarcocysts to the tawny owl, Strix aluco L., 1758 and obtained a few oocysts for a short time from its feces. However, they believed that this was an abnormal host and that another genus of strigid bird might be a better definitive host. They never attempted to name the species of the weasel sarcocysts. Original host: Neovison (syn. Mustela) vison (Schreiber, 1777), American Mink. Remarks: Foreyt and Todd (1976) collected fecal samples on 29 mink ranches (22 counties) in Wisconsin, USA, from June-December, 1975. They found that 45/79 (57%) mink feces collected from ranches that fed dry pelleted rations contained coccidian oocysts, whereas 128/244 (52%) samples from ranches that fed a wet meat mixture contained oocysts. They mentioned finding oocysts resembling an undescribed Eimeria species in mink on 19/29 (66%) of the ranches, but the number of mink with this species was not mentioned, stating only "the coccidia of domestic mink in Wisconsin are prevalent and widespread." They said the oocysts were 12.5 × 11.8 but gave no additional descriptive information. To our knowledge, this species has never been mentioned again in the literature. Original host: Neovison (syn. Mustela) vison (Schreiber, 1777), American Mink. Remarks: Ryan et al. (1982) studied skinned muskrat (Ondatra zibethica) carcasses from Connecticut and New Jersey, USA and found coccidia-like zoites in their muscles using a pepsin digestion technique. They then fed seven pairs of coccidia-free mink half of an infected muskrat carcass each. Four pairs of mink fed muskrats shed unsporulated coccidian oocysts in their feces 6-8 DPI and patency lasted 4-6 days. Sporulated oocysts were L × W (n = 80): 11.6 × 10.7 (11.5-12 × 10-11), L/W ratio: 1.1. They reported that the oocysts had a M, but we did not see one in a photomicrograph presented (their Fig. 4) , and the oocysts lacked a PG and OR. Sporocysts were L × W (n = 160): 8.8 × 6.5 (8-9 × 6-7), L/W ratio: 1.4 and had a granular SR, but lacked SB and SSB. They also fed another seven pairs of coccidia-free mink ∼200,000 sporulated oocysts each that were derived from the previous four pairs that shed the oocysts, but none of the second seven pairs shed oocysts during the course of the experiment. After 30 DPI, all 14 pairs of mink (those fed tissue cysts and those fed only oocysts) were killed and sections of gastrointestinal tract, pancreas, ileocecal lymph node, liver, kidney, lung, heart, brain, diaphragm, tongue, esophagus, and masseter muscles were examined for coccidian cysts and skeletal muscles were examined via pepsin digestion. All were negative. The authors concluded that their work was insufficient to establish whether this Hammondia-like parasite was identical to an already known species or was a new species. This interesting study certainly deserves more attention, especially now that we have molecular tools with which to work. Original host: Neovison (syn. Mustela) vison (Schreiber, 1777), American Mink. Remarks: Presidente and Karstad (1975) found meronts of a Hepatozoon in the lungs of 10/18 (56%) American minks they examined in southwestern Ontario, Canada. Meronts were found in the pulmonary parenchyma, and focal aggregations of lymphocytes, macrophages, plasma cells, and eosinophils were associated with small groups of meronts. Two kinds of meronts were seen: subspheroidal forms, 22-29 × 19-24, with a single row of 18-24 macromerozoites around the perimeter; and larger, oblong to spheroidal meronts, 29-38 × 19-24, with 34-38 micromerozoites located throughout the meront. They made no attempt to further identify or name the parasite. Original host: Neovison (syn. Mustela) vison (Schreiber, 1777), American Mink. Remarks: Sealander (1943) examined 158 mink carcasses that he obtained from southern Michigan fur-buyers in the winters of 1940 and 1941. He was looking specifically to catalog helminth parasites, but briefly mentioned that "coccidia, Isospora bigemina, were frequently noted in fecal samples." In all likelihood, he was seeing sporocysts of an unknown Sarcocystis species. Definitive host: Unknown. Intermediate host: Neovison (syn. Mustela) vison (Schreiber, 1777), American Mink. Remarks: Ramos-Vera et al. (1997) had three 2-3-month-old minks with signs of progressive neurological disease. One mink had variable numbers of sarcocysts, which measured up to 300 × 20, in multiple skeletal muscles. The sarcocyst wall had numerous elongated villar protrusions that measured 1.7 μm × 250 nm. These protrusions had microtubules and irregularly distanced minute undulations. The most important lesions were seen in the brains of all three minks. The authors said this was the first time Sarcocystis sarcocysts were described in mink muscles. The parasite was not identified to species and is considered a species inquirenda. Definitive host: Unknown. Intermediate host: Callorhinus ursinus (L., 1758), Northern Fur Seal. Remarks: Brown et al. (1974) found sarcocysts in the masseter muscle of one adolescent northern fur seal among a group of 30 pups and two adults on St. Paul Island, Pribiloff Islands, Alaska, USA. Although this was the first report of an apparent Sarcocystis species in marine mammals, the life cycle of this species is still unknown. No other information was provided. This species has not been identified since 1974 and must remain a species inquirenda. Remarks: Colegrove et al. (2011) discovered coccidian endogenous stages in the small intestine of five free-ranging California sea lions during routine postmortem examinations. In each case, multiple stages of both asexual (meronts, merozoites) and sexual (micro-and macrogamonts) stages were seen in the apical cytoplasm of distal jejunal enterocytes. Using standard histological techniques, IHC, TEM, DNA extraction, and PCR amplification of ITS-1 and 18S rRNA genes they were able to conclude that they identified previously undescribed intestinal protozoan species representing three previously uncharacterized gene sequences representing three new coccidian species that are most closely related to, but not identical with Neospora caninum. Original host: Zalophus californianus (Lesson, 1828), California Sea Lion. Remarks: Girard et al. (2016) found 16/139 (11.5%) sea lions, stranded in California, USA in 2010 and 2012, shedding oocysts in their feces. In 2011-12, they collected more fecal samples at sea lion haul-out sites at three locations in central California and 13/212 (6%) had oocysts present. Amplified ITS-1 from the samples strongly supported the presence of Coccidia A and B of Colegrove et al. (2011) . One sample produced a novel sequence that had 97%-98% pairwise similarity to previously published coccidian DNA isolates in Guadalupe fur seal tissues (Gibson et al., 2011) . Coccidia A and B were sporulated and inoculated into mice to investigate infectivity and pathogenicity, but no remarkable clinical signs or histologic changes were observed except ∼40% of the mice had slight nephritis, pneumonia, and inflammation in lung, spleen, and lymph nodes. Brain, heart, and tongue all tested negative using pan-coccidian primers that included the ITS-1 region. Results of phylogenetic analyses using ITS-1 indicated the Coccidia A, B, C, CSL Año 11, and isolates from harbor seals and Guadalupe fur seals all shared a common ancestor with N. caninum. Girard et al. (2016) concluded that additional genetic and morphologic studies are required to resolve the taxonomy of these novel marine mammal coccidia, and they believe it is likely these organisms are either a new Neospora species or a new genus in the Sarcocystidae. Girard et al. (2016) also inoculated excysted SZ of Coccidia A and B into tissue flasks of Green Monkey-derived MA104 cells. At 14, 21, and 35 DPI, 3/4 (75%) cultures had intra-and extracellular propagating organisms morphologically similar to Sarcocystis spp. Extracellular zoites were 8-11 × 1-2. As the cultures aged, the number of zoites increased suggesting they were merozoites produced via merogony, and they described active zoites to exhibit movement typical of S. neurona. Antigen slides, prepared from zoites, tested positive to S. neurona antisera from a horse and a sea lion by IFAT. PCR amplification of zoite DNA confirmed that zoites were similar to S. neurona, which allowed Girard et al. (2016) to conclude that in addition to the observed Coccidia A and B, a S. neurona-like organism is excreted by sea lions in low concentrations and sea lions are mechanical vectors. Definitive host: Unknown. Intermediate host: Zalophus californianus (Lesson, 1777), California Sea Lion. Remarks: Mense et al. (1992) did not proof the galley of their manuscript before it went to press, so their published title reads, "Acute hepatic necrosis associated with a Savmcystis-like…," instead of Sarcocystis-like. We mention this only to aid those doing a search online, who may have trouble bringing up this title. In addition, the first line of their report says, "A 10-yearold (sic) male California sea lion…" so it is not clear if the affected sea lion from an aquarium in Florida, USA was 1-or 10-years-old. This animal died after 3 days of exhibiting lethargy and anorexia. Tissues were fixed and preserved for both LM and TEM and later stained with anti-T. gondii, anti-N. caninum, and anti-S. cruzi serum; parasite stages in the liver reacted only with the anti-Sarcocystis serum. Sections of infected liver demonstrated coagulative and lytic necrosis with associated nuclear pyknosis, karyorrhexis, and hepatocyte loss in areas adjacent to parasite stages. Sarcocysts in myocytes, however, were unassociated with inflammation. Parasites in the liver divided by endopolygeny and both meronts and merozoites were seen there. Meronts were up to 32 long and contained 6-35 merozoites, each ∼5 × 1. Their TEM sections demonstrated the meronts to be located free in the hepatocyte cytoplasm, without a PV, and merozoites contained numerous micronemes, a conoid, but no rhoptries. The parasite they identified as a Sarcocystis species, likely was S. neurona, but there is no way to state that unequivocally, so their form must be relegated to a species inquirenda. GENUS CYSTOPHORA NILSSON, 1820 (MONOTYPIC) Original host: Cystophora cristata (Erxleben, 1777), Hooded Seal. Remarks: Bass et al. (2012) examined one fecal sample from a hooded seal, C. cristata, in the Gulf of Maine, USA, using a nested PCR amplification and subsequent sequencing of the 18S rRNA gene they determined the presence of Cryptosporidium sp. and found it most closely related to the genotype Cryptosporidium sp. seal 1 and seal 2 described by Santín et al. (2005) from ringed seals. Definitive host: Unknown. Intermediate host: Erignathus barbatus (Erxleben, 1777), Bearded Seal. Remarks: Bishop (1979) found a small number of sarcocysts in the skeletal muscle of the tongue of a bearded seal that had been killed by polar bears in the Arctic. Obviously, the life cycle of this species is unknown. No other information was provided. This species has not been identified since 1979 and must remain a species inquirenda. Using PCR amplification of 18S rRNA, HSP70, and COWP genes, they found two individuals (14%) positive for a Cryptosporidium sp. that represented a novel genotype they called the Weddell seal genotype. Original host: Leptonychotes weddellii (Lesson, 1826), Weddell Seal. Remarks: Dróżdż (1987) reported cherryorange, ellipsoidal oocysts with a thin, wrinkled, bilayered wall, M: absent, that measured L × W: 25.7 × 18.8 (22-31 × 14-22) , L/W ratio: 1.4, in 5/65 (8%) Weddell seals from King George Island, South Shetlands from April, 1981 to January, 1982 . No other information was provided, and this species has not been identified since 1987; thus, it must be relegated to a species inquirenda. Original host: Leptonychotes weddellii (Lesson, 1826), Weddell Seal. Remarks: Dróżdż (1987) found gray, spheroidal oocysts with a thin, smooth wall without a M, that were 25-33 wide, L/W ratio: 1.0, in 3/65 (5%) Weddell seals from King George Island, South Shetlands from April, 1981 to January, 1982 . No other information was provided, and this species has not been identified since 1987; thus, it must be relegated to a species inquirenda. Definitive host: Unknown. Intermediate host: Leptonychotes weddellii (Lesson, 1826), Weddell Seal. Remarks: Ippen and Henne (1989) examined chest, back, tongue, esophagus, diaphragm, and heart muscle tissue from a Weddell seal collected in the Antarctic. Large numbers of cysts were observed in the chest, back, tongue, and esophagus. Cross-sections of cysts were 40-70 × 50-110, and cystozoites were 4-5 × 1. Longitudinal slices of cysts were "serpentine" and measured 30-100 × 100-180. No other information was provided, and this species has not been identified since 1989; thus, it must be regarded a species inquirenda. Original host: Lobodon carcinophaga (Hombron and Jacquinot, 1842), Crabeater Seal. Remarks: Dróżdż (1987) said he found yellowgray, ovoidal oocysts with a smooth, bilayered wall without a M, that measured L × W: 30.0 (sic) × 24.3 (38-40 × 24-25), L/W ratio: 1.2, in 6/43 (14%) crabeater seals from King George Island, South Shetlands from April, 1981 to January, 1982 . No other information was provided, and this species has not been identified since 1987; thus, it must be regarded as a species inquirenda. Definitive host: Unknown. Intermediate host: Lobodon carcinophaga (Hombron and Jacquinot, 1842), Crabeater Seal. Remarks: Ippen and Henne (1989) examined chest, back, tongue, esophagus, diaphragm, and heart muscles from two crabeater seals, a male and a female, collected in the Antarctic. Male tissues were negative, but the female had a "moderate" infection with sarcocysts in her back, tongue, and esophagus. Cysts in cross-section were 40-80 × 60-110. Cysts in the esophagus measured 30-90 × 70-340. No other information was provided, and this species has not been seen again since 1989; thus, it must be relegated to a species inquirenda. Definitive host: Unknown. Intermediate host: Mirounga leonina (L., 1758), Southern Elephant Seal. Remarks: Ippen and Henne (1989) examined chest, back, tongue, esophagus, diaphragm, and heart muscles from two male elephant seals, one adult and one juvenile, collected in the Antarctic. Adult tissues were negative, but the juvenile had cysts present in the chest, tongue, and diaphragm. Cysts were smaller than those observed in Weddell and crabeater seals reported in the same paper, measuring 20-40 × 30-80 in cross-section. Ippen and Henne (1989) suggested that in the juvenile the cysts might be the result of a recent infection and had not reached their full size. No other information was given by the authors, and this species has not been reported again since 1989: thus, it must be relegated to a species inquirenda. Remarks: While examining reindeer and caribou muscle for sarcocysts in Alaska, Hadwen (1922) also found sarcocysts in the heart and esophagus of a seal. In the heart, the sarcocysts were numerous and averaged 0.43 mm × 0.17 mm. Sarcocysts in the esophagus and other muscles were larger, 0.87 mm × 0.14 mm, with the longest cyst 2.5 mm. Hadwen (1922) said these sarcocysts looked similar to those of Sarcocystis tenella of sheep. Nothing more is known about this form. Definitive host: Unknown. Intermediate host: Phoca vitulina richardii (Gray, 1864) , Pacific Harbor Seal. Remarks: Lapointe et al. (1998) reported seven Pacific harbor seals that, upon examination, had meningoencephalitis associated with S. neuronalike parasites. One seal was reported to have "rare" sarcocysts of an undetermined species in its cardiomyocytes. Original host: Pusa hispida (Schreber, 1775), Ringed Seal. Remarks: Hughes-Hanks et al. (2005) examined fecal samples of P. hispida from northern Alaska, near Barrow, using the immunofluorescent assay. Cryptosporidium was detected in 7/31 (23%) samples (5 males, 2 females). No specific identification was attempted. Original host: Pusa caspica (Gmelin, 1788) (syn. Phoca caspica), Caspian Seal. Remarks: Kuiken et al. (2006) reported a large die-off of Caspian seals in the spring and summer, 2000, which they attributed to a canine distemper epidemic, and they mentioned finding Eimeria-like oocysts in the cytoplasm of jejunal enterocytes in 1 of the of the 18 seals they necropsied. They said the oocysts "consisted of 4-8 banana-shaped zoites (2 μm × 8 μm) with a central blue nucleus and surrounded by a narrow, birefringent wall." They did not try to identify the oocysts they saw more specifically but said they were similar to those of E. phocae from har-small intestine of a young raccoon in which they also described a Cryptosporidium sp. (below). No other information was given. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Carlson and Nielsen (1982) first reported the presence of Cryptosporidium in a young male raccoon from Waterford, Connecticut, USA. The raccoon did not show any clinical signs of infection. Histology revealed endogenous stages of a Cryptosporidium sp. at the tips of the small intestinal villi, but not in the crypts. The villi were reduced in length, blunted, and increased in width; lamina propria was infiltrated with eosinophils and mononuclear cells. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Feng et al. (2007) surveyed fecal samples of wild P. lotor from the watershed of the New York City drinking water supply, USA, for Cryptosporidium spp. Using 18S rDNA and RFLP gene sequencing and analysis they reported 4/21(19%) positive samples, 1 for a cervine genotype (syn. genotype W4) and 3 for a skunk genotype (syn. genotype W13). Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Leśniańska et al. (2016) collected feces from P. lotor that were introduced and are now invasive, to Europe; 32 samples were from two areas in Poland (Kostrzyn on the Oder, and Warta Mouth National Park), and 17 samples from Germany (Müritz National Park, Mecklenburg-Vorpommern). Cryptosporidium was detected in 17/49 (35%) samples (14 from Poland, 3 from Germany). Amplification of 18S rRNA, COWP, and actin genes demonstrated that 9/17 (53%) infected raccoons had Cryptosporidium skunk genotype. This study was the first evidence of Cryptosporidium in raccoons from Poland and Germany. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Martin and Zeidner (1992) described a case of cryptosporidiosis in a juvenile P. lotor found moribund in Fort Collins, Colorado, USA. The animal was emaciated, dehydrated, and had diarrhea. Microscopically, Cryptosporidium endogenous stages, spheroidal to ovoidal, 2-7 wide, were seen on intact intestinal villi. The raccoon also was infected with coronavirus and parvovirus. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Zhou et al. (2004) collected the feces of 471 mammals from four counties in the Chesapeake Bay area of Maryland, USA; they found 2/51 (4%) raccoons (presumably P. lotor) to be infected with a Cryptosporidium skunk genotype. The species and genotype of Cryptosporidium in each fecal sample was determined by a PCR-RFLP method based on the 18S rRNA gene. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Ziegler et al. (2007) collected fecal samples of wild P. lotor in the New York City Watershed, southeastern New York state, USA. Samples were examined by LM and by polyclonal Cryptosporidium antigen-capture ELISA (considered positive based on an optical density ≥0.050). They found Cryptosporidium in 49/173 (28%) samples. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Robel et al. (1989) collected raccoons from two locations in Kansas, USA and reported that 8/36 (22%) raccoons from Ft. Riley and 25/92 (27%) from rural Manhattan had eimerian oocysts in their rectal contents. However, because all intestinal tracts had been frozen for 1-3 months at −5°C, which should have prevented sporulation, it is not clear to us how they were able to state these were eimerian oocysts. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Snyder (1984 , 1988 reported oocysts of Eimeria spp. in 67/100 (67%) fecal samples from P. lotor in Illinois, USA. He examined the samples for C. parvum, using sucrose flotation (1984) or an indirect immunofluorescent detection procedure (1988) and reported that "Eimeria spp. oocysts were routinely seen in samples but never exhibited any fluorescence." Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Foster et al. (2004) recovered oocysts of three Eimeria spp. from raccoons collected in Key Largo, Monroe county, Florida, USA; two were previously described (E. procyonis, E. nuttalli), but the third form, found in 2/61 (3%) samples, was unknown. Oocysts were ellipsoidal, L × W (n = 22): 29.2 × 15.7 (28-31 × 14-17), L/W ratio: 1.9; with a smooth, 2-layered wall; M, OR, PG: all absent. Sporocysts were ellipsoidal, 10.1 × 7.7 (10-11 × 6-8); SB, SSB, PSB, SR: all absent. Foster et al. (2004) speculated this was likely a spurious coccidium originating from a food item passing through the raccoon's gastrointestinal tract. Definitive host: Procyon lotor (L., 1758), Raccoon. Intermediate host: Unknown. Remarks: Adams et al. (1981) mentioned finding two naturally-infected raccoons in Illinois, USA that were discharging Sarcocystis-like sporocysts in their feces, but they did not attempt to determine the species. Sporocysts were ellipsoidal or had one side flattened, L × W (n = 17): 13.0 × 9.3 (11-13 × 8-10), with a smooth wall; SB, SSB, PSB: all absent; SR: present, and SZ were "elongate" lying lengthwise in SP. An attempt to infect four pigs resulted in no sarcocysts in pig muscles or other tissues at necropsy and no oocysts in the pig's feces. Remarks: Dubey et al. (1990b) received 45 raccoons from Ohio, USA for experiments to develop an oral rabies recombinant vaccine for wildlife. One juvenile female was unsteady on its feet, had difficulty with balance, and its head was constantly turned to the left. Seven days after arrival, it was anorectic, had mucopurulent nasal and serous ocular discharge, and was moribund. On necropsy they observed the carcass was dehydrated, mucous membranes were pale, and lungs were consolidated. Numerous developmental stages of a Sarcocystis species were found in the cytoplasm of macrophages, neurons, and multinucleated giant cells. In mature meronts, merozoites were arranged in a rosette around a residual body; individual merozoites measured 4-5 × 1 and were seen in mononuclear cells located within meningeal blood vessels. Organisms were located in host cell cytoplasm without a PV, and merozoites had no rhoptries. Moderate numbers of sarcocysts were found in striated muscles of the heart, tongue, diaphragm, esophagus, masseter, and extraorbital muscles. In all likelihood, this species is S. neurona but must be considered a species inquirenda at this time. Definitive host: Procyon lotor (L., 1758), Raccoon. Intermediate host: Unknown. Remarks: Foster et al. (2004) collected sporocysts from the feces of 2/61 (3%) raccoons on Key Largo, Monroe county, Florida, USA. Sporocysts were ellipsoidal, L × W (n = 10): 13.3 × 8.5 (12-14 × 7-10). There was no attempt to identify the species. Definitive host: Procyon lotor (L., 1758), Raccoon. Intermediate host: Unknown. Remarks: Robel et al. (1989) collected raccoons from two locations in Kansas, USA and reported that 5/92 (5%) from rural Manhattan, but 0/36 from Ft. Riley, had sporocysts of a Sarcocystis sp. in their rectal contents. However, these numbers only were present in their Table I , and they never mentioned the presence of Sarcocystis in the body of their paper. It also should be noted that all intestinal tracts had been frozen, for 1-3 months at −5°C, before they were examined, so it is not clear to us if the sporocysts they saw had SZ in them or not. Definitive host: Unknown. Intermediate host: Procyon lotor (L., 1758), Raccoon. Remarks: Seneviratna et al. (1975) examined the muscles of domestic animals and a few wild animals killed in Detroit, Michigan, USA from April to September, 1973. They found 2/6 (33%) raccoons, both female, with sarcocysts in their muscles. They were unable to make a species identification; thus, another species inquirenda. Definitive host: Procyon lotor (L., 1758), Raccoon. Intermediate host: Procyon lotor (L., 1758), Raccoon. Remarks: Thulin et al. (1992) reported an adult male raccoon with protozoal encephalitis associated with a Sarcocystis-like organism and concurrently infected with canine distemper. The Sarcocystis species was intracellular in the cytoplasm, without a PV. Histologically, sarcocysts were present in skeletal muscle and occasionally in the heart; these cysts were 19 × 25 to 40 × 506 and had striated walls that were 1.5-2.0 thick. Individual merozoites were 2.0 × 1.5. A second raccoon had numerous Sarcocystis-type oocysts in the lamina propria of its small intestine. It is possible that the form in the first raccoon was S. neurona because of its intracellular location without a PV, with the raccoon serving as an intermediate host; similarly, the isosporoid oocysts in the gut of the second raccoon may represent another species of Sarcocystis, with the raccoon serving as the definitive host. Both forms can only be considered species inquirendae at this time. No clinical signs were observed in the pandas at the time of feces collections. One 18-year-old panda was positive for Cryptosporidium oocysts that were L × W (n = 50): 4.6 × 4.0 (4-6 × 3-5). Using the sequences of 18S rRNA, HSP70, COWP, and actin genes, Liu et al. (2013) discovered that this parasite represented a new genotype of Cryptosporidium, which was most closely related to the Cryptosporidium bear genotype, and they designated this new genotype, Cryptosporidium giant panda genotype, but did not give it a binomial. Original host: Helarctos malayanus (Raffles, 1821), Sun Bear. Remarks: Siam et al. (1994) examined 81 captive carnivores in the Giza Zoological Gardens Zoo, Egypt, including two sun bears; they found 1/2 (50%) sun bears to be passing oocysts of a Cryptosporidium sp. No other information was provided. Original host: Helarctos malayanus (Raffles, 1821), Sun Bear. Remarks: Wang and Liew (1990) first reported the presence of oocysts of Cryptosporidium in both captive sun bears in a bird park in Taiwan. Parasites were detected in fecal smears stained with modified Ziehl-Neelsen karbolfuchsin staining method. No other information was provided. Definitive host: Unknown. Intermediate host: Helarctos malayanus (Raffles, 1821), Sun Bear. Remarks: Latif et al. (2010) examined one Malayan sun bear in a zoo in Korea and found that it had sarcocysts of Sarcocystis only in the diaphragm, but they did not find sarcocysts in the tongue, esophagus, heart, or skeletal muscles. There was no attempt to identify the species beyond genus so their finding can only be considered a species inquirenda. (4 SPECIES) Original host: Ursus arctos L., 1758 (syn. Ursus horribilis Ord, 1815), Brown Bear. Remarks: Gau et al. (1999) reported the presence of "gastrointestinal coccidia" in 8/56 (14%) fecal samples of U. arctos from the Central Canadian Arctic (Northwest Territories, Canada), but they neither measured oocysts nor identified the genus of the coccidia they observed (probably because fecal samples all had been frozen in the field and stored at −20°C until analyzed, which would have prevented sporulation). They suggested that the coccidians may be enzootic in U. arctos in the central Arctic, rather than incidentally occurring through the ingestion of infected prey species. This species, or these species, can only be considered species inquirendae. Original host: Ursus arctos L., 1758 (syn. Ursus horribilis Ord, 1815), Brown Bear. Remarks: Bemrick and O'Leary (1979) reported that a 1-week-old female grizzly (brown) bear cub, from the Como Park Zoo, St. Paul, Minnesota, USA had severe diarrhea. The cub weighed only 454 g and died. Routine histological examination of the lamina propria of the small intestine showed exceptionally large asexual stages (meronts) present, but these were not measured and, although four photomicrographs of tissue sections were published, no structural measurements were given. Bemrick and O'Leary (1979) mentioned that fecal samples of other grizzly bears in the zoo had been examined periodically in previous years, but no coccidian oocysts were found. They could not say with certainty that these endogenous stages were the etiological agent of the diarrhea seen in the bear cub or not. Their coccidium must be considered a species inquirenda. Original host: Ursus arctos L., 1758 (syn. Ursus horribilis Ord, 1815), Brown Bear. Remarks: Siam et al. (1994) examined 81 captive carnivores in the Giza Zoological Gardens Zoo, Egypt, including brown bears; they found 2/7 (29%) brown bears passing oocysts of a Cryptosporidium sp. No other information was provided. Original host: Ursus maritimus Phipps, 1774, Polar Bear. Remarks: Siam et al. (1994) examined 81 captive carnivores in the Giza Zoological Gardens Zoo, Egypt, including polar bears; they found 1/7 (14%) polar bears passing oocysts of a Cryptosporidium sp. No other information was provided. Original host: Ursus arctos L., 1758 (syn. Ursus horribilis Ord, 1815), Brown Bear. Remarks: Aghazadeh et al. (2015) examined fecal samples from European brown bears collected in Croatia and found 1/94 (1%) positive for Eimeria oocysts. The authors provided no description except that oocysts were unsporulated and had a thin, white-gray double oocyst wall, but no M. They noted that the oocysts could have occurred incidentally via infected prey. Original host: Ursus arctos L., 1758 (syn. Ursus horribilis Ord, 1815), Brown Bear. Remarks: Isospora oocysts were found in 2/7 (29%) brown bears in the Leningrad Zoo, former USSR. The animals were imported from north of the former USSR, which likely is the natural habitat for both the host and these oocysts. Oocysts sporulated in 4 days in 2% K 2 Cr 2 O 7 solution at 24°C. The oocysts had two forms, with a wall ∼1.5 thick. Spheroidal oocysts were 24.6 (19-32), whereas ovoidal oocysts were L × W: 26.4 × 24.0 (23-33 × 21-29), L/W ratio: 1.1; M: absent; OR: present (line drawing); PG: present. Ovoidal to piriform sporocysts were L × W: 16.9 × 11.6 (15-19 × 11-13), L/W ratio, 1.5; SB, SSB: both present; PSB: absent; SR: present; SR characteristics: a clump of granules or granules spread among SZ; SZ: crescent-shaped. Yakimoff and Matschoulsky (1935) thought this was not a parasite of bears, but may be a bird isosporan because its morphological features were similar to sporulated oocysts described as Isospora lacazei (Labbé, 1893) . Its occurrence in this bear's feces may be explained by the fact that bears lick their paws contaminated by the soil where the sparrows defecate. No one knows for sure. Definitive host: Unknown. Intermediate host: Ursus americanus Pallas, 1780, American Black Bear. Remarks: Crum et al. (1978) found 6/53 (11%) U. americanus collected from six states in the southeastern United States to have sarcocysts, which they found histologically in the cardiac and skeletal muscles. No other information was given. Definitive host: Unknown. Intermediate host: Ursus americanus Pallas, 1780, American Black Bear. Remarks: Sarcocysts were found in 1/374 (∼0.3%) black bears legally shot in Pennsylvania, USA, but only one cyst was found in this bear. The sarcocyst from this bear was structurally different from the sarcocysts found in two bears, during the same collection, which allowed Dubey et al. (2008a) to name a new species at that time, S. ursusi. The wall of these sarcocysts was ∼2 thick and had finger-like villi on the cyst wall giving the sarcocyst wall a striated appearance that the sarcocysts of S. ursusi lacked. Nothing else is yet known about the identity of the Sarcocystis species in bears with these kinds of sarcocysts. Definitive host: Unknown. Intermediate host: Ursus americanus Pallas, 1780, American Black Bear. Remarks: Dubey et al. (1998b) were able to examine diaphragm, abdominal muscles, and carcass muscles from 92 hunter-killed bears in North Carolina, USA. They found two sarcocysts in cross-sections of muscle from one bear (1%). The sarcocysts were 45 × 37.5 and 67.5 × 50. Under TEM they saw that the bradyzoites butted against the ground substance of the cyst wall; five longitudinally-cut bradyzoites were 6.0-6.6 × 2.5-3.3 and contained organelles typically found in Sarcocystis bradyzoites. Dubey et al. (1998b) reiterated a previous (perhaps questionable) argument that the structure of the sarcocyst wall is a reliable taxonomic criterion to distinguish Sarcocystis species within a given host. They concluded their reasoning as to why they did not call this the same species as the one seen by Zeman et al. (1993) , below, by saying, "A Sarcocystis-like parasite has been reported as causing fatal hepatitis in a black bear from South Dakota (Zeman et al., 1993) and in two polar bears from Alaska (Garner et al., 1997) . Only schizonts and merozoites were found in bears that died of hepatitis; sarcocysts were not seen." Thus, the observational data between this report and those of Zeman et al. (1993) and Garner et al. (1997) is not comparable, and all of these can only be considered as species inquirendae. Definitive host: Unknown. Intermediate host: Ursus maritimus Phipps, 1774, Polar Bear. Remarks: Garner et al. (1997) diagnosed fatal hepatic sarcocystosis in two zoo polar bears in Anchorage, Alaska, USA. Gross lesions were icterus and systemic petechiae, whereas microscopic lesions were detected only in the liver and included severe random necrotizing hepatitis with hemorrhage. They observed only asexual stages of the parasite within hepatocytes along with rare extracellular zoites. This parasite divided by endopolygeny, and occasionally merozoites formed rosettes around a central residual body. TEM of merozoites showed a conoid and a small number of micronemes at their apical pole, a central N, but rhoptries were absent. These parasites failed to react with anti-Neurospora sp., anti-T. gondii, and anti-S. neurona sera. Nothing more about this parasite is known, including the life cycle. Thus, it must be considered a species inquirenda. Definitive host: Unknown. Intermediate host: Ursus americanus Pallas, 1780, American Black Bear. Remarks: Zeman et al. (1993) reported a case of fatal sarcocystosis in an American black bear from a wild-animal park, Black Hills, South Dakota, USA; initially, they found different developmental stages of a protozoan parasite in the cytoplasm of several hepatocytes. There was no inflammation associated with parasitized hepatocytes, but they saw both inflammation and necrosis associated with maturation and rupture of mature meronts. Zeman et al. (1993) reported that mature meronts occupied 50%-80% of hepatocyte cytoplasm and moved the HCN to the periphery of the cell. They also noted that the meronts divided by endopolygeny, characteristic of Sarcocystis species. Mature meronts were 30 × 20 and contained up to 36 merozoites. When liver tissue sections were deparaffinized, they reacted with antiserum to T. gondii, N. caninum, and S. cruzi, but the parasite tissue only reacted with S. cruzi, not with T. gondii or N. caninum antisera. Zeman et al. (1993) compared the hepatic lesions seen in the bear to those reported from a sea lion (Mense et al., 1992) and said these lesions also were identical to those described in a 2-day-old dog with congenital sarcocystosis (Dubey et al., 1992a) . The canine parasite was morphologically and antigenically identical to Sarcocystis canis, but unlike in the bear, the S. canis infection was found in many of the dog's tissues including skin, brain, liver, and lungs. This was the first report of a fatal hepatic sarcocystosis in a bear, but its true identity is still unknown. were detected in 87/1,294 (7%) cats of all age categories but mostly in those <6-months-old. Oocysts found were not determined to genus or species, and the authors admit that everything was reported as Isospora sp. or as "coccidia." Original host: Felis catus L., 1758, Domestic Cat. Remarks: McGlade et al. (2003) examined 418 fecal samples of 125 domestic cats, from various sources in Perth, Western Australia; 40 randomly-selected samples were screened by nested PCR of the 18S rRNA gene. None of the 418 samples was positive by microscopy, but 4/40 (10%) PCR-sequenced samples were positive for Cryptosporidium spp. and one successfully-sequenced PCR product, "most closely resembled" C. baileyi (commonly found in chickens). Original host: Felis catus L., 1758, Domestic Cat. Remarks: Nash et al. (1993) studied fecal samples of cats from 8 farms in Glasgow, Scotland, by several tests including a specific monoclonal antibody against Cryptosporidium (Northumbria Biologicals); Cryptosporidium was detected in 7/57 (12%) cats (3 kittens, 4 adults) on three farms. They euthanized 32 cats, all without diarrhea, and processed them for histology. Histology revealed the presence of endogenous developmental stages and oocysts in the epithelium of the ileum, and two types of oocysts were recorded, smaller, 5.0 × 4.5, and larger, 6.0 × 5.0, which were detected in one farm, whereas in the other two farms only smaller oocysts were found. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Ogassawara et al. (1986) surveyed the feces of domestic cats from different areas in São Paulo, Brazil, and reported finding oocysts of a "species" they called C. curyi in 8/215 (4%) cats. However, the citation they gave for this species (Ogassawara, S., Benassi, S., Larsson, C.E., Hagiwara, M.K. 1986 . Cryptosporidium curyi n. sp. in the feces of Felis catus domesticus in the city of São Paulo, Brazil. Rev. Microbiol., São Paulo.) apparently was never published. They refer to C. curyi, and the paper in which it was presumably published as follows: "This agent has recently been described by us and is still being studied for its true identity." This species and citation do not appear in Wikipedia, Google Scholar, or in more recent reviews of Cryptosporidium species names by Plutzer and Karanis (2009) Original host: Felis catus L., 1758, Domestic Cat. Remarks: Asahi et al. (1991) reported a cat in Japan, naturally-infected with "a small type of Cryptosporidium sp. oocysts," measuring 4.5 wide. Necropsy showed endogenous stages in the villous epithelia of the small intestine and cecum, but not in the stomach. Oocysts from this cat were inoculated per os into six cats (weight < 1 kg), and all six discharged oocysts; the prepatent period was 8-10 days and peak oocyst shedding was 14-19 DPI, followed by 7-14 more peaks during patency (i.e., 69-203 DPI). All the cats were asymptomatic. After patency ended, the cats were injected s.c. with prednisolone, 10 mg/kg daily for 4-9 days, which caused the recurrence of oocyst shedding. Experimental infections with oocysts in specific pathogen-free (SPF) mice, BALB/c mice, BALB/c mice injected with hydrocortisone acetate, SPF Wistar rats, SPF guinea pigs, and dogs, all failed. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Barr et al. (1994a) diagnosed cryptosporidiosis in three cats suffering from diarrhea, based on the presence of Cryptosporidium oocysts in their feces. The cats were administered paromomycin for 5 days, and the diarrhea resolved 5 days after the last dose in two cats, and within 30 days in the third cat. No other information is given. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Barr et al. (1994b) diagnosed Cryptosporidium oocysts and Toxocara cati eggs in the feces of a 6-month-old spayed domestic cat that had diarrhea for 2 months. The cat was properly vaccinated and was both FeLV and FIV negative. It was administered pyrantel pamoate (against Toxocara), metronidazole (against protozoans), and a special diet (Hill's c/d), but after 14 days the cat still had diarrhea and still shed Cryptosporidium oocysts in its feces. It was then given paromomycin, 165 mg/kg, p.o., twice daily for 5 days, and Cryptosporidium was not detected in its feces 1, 8, or 34 days after treatment. On days 1 and 8, the diarrhea persisted, but by day 34 the cat was nondiarrheic ∼80% of the time. Thus, paromomycin successfully treated the Cryptosporidium infection, although it was not identified to species. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Borkataki et al. (2013) surveyed fecal samples of stray cats in Jammu, a humid subtropical zone in northwestern India. Oocysts of Cryptosporidium were detected in 4/100 (4%) samples, but not identified to species. and Cryptosporidium oocysts were detected in 2/51 (4%) cats using a malachite green negative stain, and in 3/51 (6%) cats with ELISA. Positive samples mostly originated from young animals and did not depend on the sex or breed. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Cox et al. (2005) examined fecal samples of various domestic, native, and feral animals during a survey of protozoal, bacterial, and viral pathogens, in four drinking-water watersheds in Sydney, Australia. Fecal samples were fluorescence-stained and examined with an epifluorescence microscope. Oocysts of Cryptosporidium were detected in 3/7 (43%) domestic cats, but not in the only feral cat examined. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Dado et al. (2012) examined 625 soil samples and 79 fecal samples ("presumably from dogs and cats") from 67 playgrounds and public parks in southeastern Madrid, Spain. Soil samples were examined using the modified Telemann and MIF methods and fecal samples by a modified Ziehl-Neelsen staining, followed by a rapid immunochromatographic assay. Cryptosporidium oocysts were only found in 7/79 (9%) fecal samples found in four parks. Original host: Felis catus L., 1758, Domestic Cat. Remarks: de Oliveira Lemos et al. (2012) examined fecal samples of diarrheic (acute or chronic) domestic cats from Rio de Janeiro, Niterói, and Praia de Mauá, Brazil. Cryptosporidium oocysts were detected in 5/60 (8%) samples, and 4 of the positive samples also were FeLV-positive. They noted that animals infected with retroviruses were more prone to the Cryptosporidium infection and may exhibit more severe clinical signs. Original host: Felis chaus Schreber, 1777, Jungle Cat. Remarks: Dubey and Pande (1963b) found a "sporulated stage" that measured 11 × 7 (10-12 × 7-8), L/W ratio: 1.2-1.5; this "stage" had 4 SZ that were 7-9 × 1-2, each with a large RB at their broader end. They called these stages "Cryptosporodial/Coccidial bodies," in their discussion and labeled their drawing ( Fig. 16) a Cryptosporidium. It's likely they were looking at a sporocyst of a Sarcocystis sp. because we think the cyst was too large to be a Cryptosporidium. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Egger et al. (1990) described intestinal cryptosporidiosis in an 8-year-old boy who suffered from coughing and bronchitis prior to having watery, nonbloody diarrhea, vomiting, and colics. The boy became ill during a visit to a dairy farm, where some kittens suffered from diarrhea and showed failure to thrive. The boy's stool was negative for rotaviruses and pathogenic bacteria, but using auramine fluorescence staining, oocysts of Cryptosporidium were found both in the stool of the boy and in feces of a skinny kitten he had contact with. This indicated that the infection "was acquired from a cat that had probably been infected by feces from calves." Original host: Felis catus L., 1758, Domestic Cat. Remarks: Hoopes et al. (2015) examined fecal samples of rural, free-roaming, and pet cats from urban areas of Saskatchewan and from a rural region in southwestern Alberta, Canada. Cryptosporidium was detected in 11/161 (7%) free-ranging cats, only in Saskatchewan, but not in 31 cats from pet shops in Saskatchewan nor in 27 cats tested from rural Alberta. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Huber et al. (2002) examined fecal samples of 28 shelter, and 20 household adult cats in Rio de Janeiro, Brazil; all cats were apparently healthy. Oocysts of Cryptosporidium were detected in 6/48 (12.5%) samples, 5/20 (25%) household cats, and 1/28 (4%) shelter cats. The authors said that household cats were more parasitized with protozoa, whereas more shelter cats harbored worms. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Korkmaz et al. (2016) examined fecal samples of pet and stray cats in Kirikkale province, Turkey. Oocysts of Cryptosporidium were detected in 1/100 (1%) samples. Original host: Felis chaus Schreber, 1777, Jungle Cat. Remarks: Lim et al. (2008) found Cryptosporidium oocysts in the feces of 1/3 (33%) jungle cats examined at Zoo Negra, Kuala Lumpur, Malaysia. No other information was given. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Lucio-Forster and Bowman (2011) examined 1,272 fecal samples from two shelters and 50 from foster homes in upstate New York, USA. Cryptosporidium oocysts were detected in 50/1,322 (4%) samples. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Marks et al. (2004) examined 416 feces of 104 domestic shorthair kittens, 8-16-weeksold, in Davis, California, USA, which were naturally-exposed to Cryptosporidium spp. The kittens were housed individually, and their fecal samples were collected once daily for four consecutive days. Samples were examined by five diagnostic tests to evaluate and compare their sensitivity: (1) a modified Ziehl-Neelsen acid-fast staining technique (mZN); (2) direct immunofluorescence (DIC) by a commercial kit (Merifluor Cryptosporidium/Giardia direct immunofluorescent kit) that used a monoclonal antibody against antigen of C. parvum; (3) enzyme immunoassay-1 (EIA-1) that used a monoclonal antibody (Premier Cryptosporidium EIA); (4) EIA-2 that used a polyclonal antibody (ProSpecT Cryptosporidium microplate assay) and (5) EIA-3 that used a polyclonal antibody (ProSpecT Cryptosporidium rapid assay) to detect Cryptosporidium antigen in feces. EIA-3. Marks et al. (2004) suggested higher sensitivity of mZN over DIC due to the preparation procedure in the manufacturer's guidelines (loss of oocysts during the rinsing procedure), and because oocyst fluorescence intensity was variable. A markedly low sensitivity of EIA-3 was likely caused by its high detection limit compared with that of EIA-1 and EIA-2. To summarize: EIA-2 and EIA-1 had the highest sensitivities when only a single fecal sample was examined, and mZN and EIA-1 had similar sensitivities when two consecutive samples were examined. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Mekaru et al. (2007) , in northern California, USA, examined feces of 344 diarrheic and nondiarrheic cats from four animal shelters. Samples were examined by three methods to evaluate/compare their specificity and sensitivity: flotation, commercial available enzyme (ProSpecT Cryptosporidium Microplate Assay), and nonenzymatic (ImmunoCard STAT! Cryptosporidium/Giardia Rapid Assay, Xpect Giardia/Cryptosporidium) immunoassay; these were compared with a reference standard, the MeriFluor direct immunofluorescence assay. They wanted to test the credibility of using human-based immunoassays for the diagnosis of Cryptosporidium spp. infections in cats and other animals. Cryptosporidium spp. was detected in 14/344 (5%) cats (9/177 diarrheic, 5/177 nondiarrheic). Two diarrheic cats were coinfected with Giardia. Flotation only detected 3/14 positive samples; the best specificity and sensitivity was found using the enzyme immunoassay ProSpecT Cryptosporidium Microplate Assay. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Mirzaghavami et al. (2016) surveyed fecal samples of stray cats in Tehran, Iran. Cryptosporidium oocysts, measuring 6.9 × 4.6, were detected in 5/50 (10%) samples. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Nutter et al. (2004) surveyed the blood and fecal samples of feral cats and healthy pet cats (mostly strays or in shelters) in rural Randolph county, North Carolina, USA. Fecal samples were concentrated by sedimentation, and then tested by a commercially-available indirect fluorescent antibody test (Merifluor, Meridian Diagnostics, Cincinnati, Ohio) . Cryptosporidium was detected in 6/87 (7%) feral and in 4/66 (6%) pet cats. The antibody test reacted with both C. parvum and C. felis and, thus, was not able to distinguish between these Cryptosporidium species. Original hosts: Bos taurus L., 1758, Aurochs; Felis catus L., 1758, Domestic Cat. Remarks: Pavlásek (1983) isolated oocysts of Cryptosporidium from a 12-day-old calf in the Czech Republic, purified them on a sugar gradient, and inoculated 5 × 10 5 sporulated oocysts per os into a 21-day-old coccidia-free cat and two chickens, and examined their feces for 45 DPI. The cat started to discharge cryptosporidial oocysts 5-12 DPI and diarrhea was observed 5-9 DPI. Chickens discharged oocysts within several hours after the inoculation until 2 DPI, so the author supposed it was just a passage through the gut and that their transmission was not successful. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Poonacha and Pippin (1982) reported a case of a 5-year-old male domestic longhair cat suffering from anorexia, weight loss, and persistent diarrhea for 2.5 months. The cat was FeLV negative, and did not respond to treatment with Azithromycin, B-12, B-plex, fluids, and lincocin. Both fecal flotation and direct microscopic examination of feces also were negative. Exploratory surgery followed by necropsy revealed dilatation and thickening of the small intestine and cecum, enlarged mesenteric lymph nodes, fusion of small intestinal villi, increased number of goblet cells, and hyperplastic crypt epithelium with a few dilated cystic crypts containing necrotic cell debris. Changes were apparent only in the small intestine and cecum, where numerous round or ovoidal organisms, 1-3 wide, were found located more in crypts and the lower half of the villi, and only rarely on the villous tips. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Tzipori and Campbell (1981) did a serosurvey for antibodies to Cryptosporidium by an indirect immunofluorescence test of the sera of cats, probably all from the Glasgow area, United Kingdom; they found 20/23 (87%) sera were positive. This was the first serological procedure for the detection of antibodies against Cryptosporidium, but there was no attempt to identify the species. (29), a breeding establishment (10), and private homes (127); none of the cats showed any clinical symptoms. Samples were examined by nested PCR amplification and sequencing of the 18S rRNA gene. Cryptosporidium was detected in 34/345 (10%) samples; five were identified as Cryptosporidium rat genotype III, and one as Cryptosporidium related to C. rat genotype III. This was the first report of Cryptosporidium rat genotype III in cats. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Between 1997 and 2007 , Gates and Nolan (2009 surveyed fecal samples of cats at the University of Pennsylvania, USA; "Cystoisospora rivolta-like" oocysts were detected in 19/1,566 (1%) samples, mostly in cats less than 3-years-old but increased when the cats were >13-years-old. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: During a survey on endoparasites, Barutzki and Schaper (2003) Original host: Felis catus L., 1758, Domestic Cat. Remarks: Christie et al. (1976) surveyed feces from cats, all >3-months-old, from a humane shelter in Ohio, USA; they found 4/1,000 (0.4%) samples to have oocysts of an Eimeria species. After sporulation in 2.5% K 2 Cr 2 O 7 solution, they inoculated ∼1,000 oocysts into 16 mice and 2 cats. After 1 month, the mice were fed to 4 SPF cats. None of the 6 cats shed oocysts within the next 30 days. Thus, this must be considered a species inquirenda. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Milstein and Goldsmid (1997) Original host: Felis catus L., 1758, Domestic Cat. Remarks: Epe et al. (1993) , in Germany, surveyed fecal samples from cats between 1984 and 1991 and found 53/1,147 (5%) to pass Isospora spp. oocysts, which were not identified to species. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Hoopes et al. (2015) examined fecal samples of rural, free-roaming, and pet cats from urban areas of Saskatchewan, and from a rural region in southwestern Alberta, Canada. Isospora spp. were detected in 19% rural cats from Alberta, 6% free-ranging cats in Saskatchewan, and in 3% pet cats in Saskatchewan. Moreover, blood was observed in two samples to be positive for Isospora. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Korkmaz et al. (2016) examined fecal samples of owned and stray cats in Kirikkale province, Turkey; oocysts of Isospora spp., which were not identified to species, were detected in 31/100 (31%) samples. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Malloy and Embil (1978) examined fecal samples of stray cats in Halifax, Nova Scotia, Canada. Oocysts of Isospora spp. were detected in 19/299 (6%) cats, mostly in young animals 0.5-2-years-old, but no attempt was made to identify the species of these isosporans. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Bos taurus L., 1758, Aurochs. Remarks: Böttner et al. (1987) surveyed muscle samples by LM, TEM, and SEM of 500 beef cattle slaughtered in New Zealand and found 100% to be infected; of these, 399/500 (80%) were infected with thick-walled sarcocysts, but they were not able to distinguish whether these sarcocysts were S. hirsuta or S. hominis. They took 305 cysts from 31 samples of diaphragm and esophagus, measured them, and examined them via LM, TEM, and SEM; cyst wall thickness was ∼5 (2-7) and was normally distributed throughout the cyst. Protrusion widths were ∼1.0 (0.6-2.2) and they formed a continuous, though skewed, distribution. They fed these thick-walled cysts to 3 cats and one human volunteer; the human did not pass sporocysts, but 1/3 (33%) cats shed a few sporocysts in its feces on 11-17 DPI. Nonetheless, they chose not to name this form so it must remain a species inquirenda, until further study. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Christie et al. (1976) surveyed feces from cats, all >3-months-old, from a shelter in Ohio, USA, but found only 2/1,000 (0.2%) samples had sporocysts of a Sarcocystis species. These sporocysts were orally inoculated into mice on the same day they were collected from cats. The mice were killed 1-6 months PI, and unstained squashes of skeletal muscles and brain were examined for cysts of Sarcocystis species and Toxoplasma. The remainder of mouse carcasses were homogenized in a blender and force-fed to 29 SPF cats. Neither of the two mice that had sarcocysts were infectious for either mice or cats. This indicated the species was not S. muris. Synonym: Cryptosporodial/Coccidial (sic) bodies of Dubey and Pande, 1963b. Definitive host: Felis chaus Schreber, 1777, Jungle Cat. Intermediate host: Unknown. Remarks: Dubey and Pande (1963b) looked at the feces of three F. chaus and found coccidian oocysts in two. They said these oocysts represented six species. Three of the forms they identified were already named, E. cati Yakimoff, 1933 , E. felina Nieschulz, 1924b , and I. rivolta (Grassi, 1879 ) Wenyon, 1923 , and two of the forms they named as new species, E. hammondi and E. mathurai. The sixth form they found was thought to represent a species of Cryptosporidium, but these "oocysts," of course were sporocysts of a Sarcocystis species, and Levine and Ivens (1981) corrected the name. The sporocysts were ellipsoidal and measured L × W: 11 × 7 (10-12 × 7-8), L/W ratio, 1.6; the sausage-shaped SZ measured 7-9 × 1-2 and had a clear RB at their rounded ends. The presence of an SR was not mentioned but their drawing (their Fig. 16 ) indicated it consisted of large globules congregated at one end of the SP. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Bos taurus L., 1758, Aurochs. Remarks: Dubey and Streitel (1976c) did crossinfections between dogs, cats, sheep, pigs, and cattle from Iowa or Ohio, USA. Cats fed bovine tissues shed sporocysts in small numbers; two cats had patent periods that lasted for 2 and 12 days after ingesting bovine tissue; sporocysts (n = 72) were 12.4 × 8.5 (11-14 × 8-11). The sporocysts they measured in cat feces were smaller than those in canine feces after ingesting infected bovine flesh. Fayer et al. (1976) earlier the same year fed 16 cats Sarcocystis-infected musculature from bovines, but none of them became infected. Dubey and Streitel (1976c) suggested that separate species of Sarcocystis parasitize cattle in different areas of the United States, but they did not suggest a name for any of their Sarcocystis species. Definitive host: Unknown. Intermediate host: Felis catus L., 1758, Domestic Cat. Remarks: Edwards et al. (1988) reported numerous cysts in a 1.5-year-old cat from College Station, Texas, USA. Sarcocysts were found in muscles of the heart, forelimbs, hind limbs, diaphragm, eyes, and intercostal spaces. Sarcocysts were (n = 50): 48 (30-60) wide × 1.5-2.2 long, had a thick wall, and septa subdivided the cyst into compartments, all with banana-shaped bradyzoites that were 7-10 long. TEM showed the primary cyst wall differed slightly from those of sarcocysts found in cats by Kirkpatrick et al. (1986) , and now thought to be identical with S. felis (Dubey et al., 1992b) . Primary cyst walls seen by Edwards et al. (1988) had a Pvm with regularly-spaced, villus projections, which were round in cross-section and all about 1 long, whereas the villus projections of Kirkpatrick et al. (1986) and Dubey et al. (1992b) had shorter, irregularly-spaced projections. This led Edwards et al. (1988) to conclude that the form they saw may be a different species. No inflammatory response was associated with the sarcocysts in any of the muscles examined. This form must also be relegated to a species inquirenda. Definitive host: Unknown. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Nanger granti (Brooke, 1872) (syn. Gazella granti), Grant's Gazelle. Remarks: Janitschke et al. (1976) found sarcocysts in Grant's gazelle in Tanzania, fed infected flesh to both cats and dogs, and reported finding sporocysts and sporulated oocysts in the feces of both. Sporocysts shed by the cat were L × W: 13 × 9 (11-16 × 8-12), L/W ratio, 1.4; the prepatent period in cats was 20 days and patency lasted 44-48 days. Janitschke et al. (1976) thought they were dealing with two Sarcocystis species, but without further infection and/or molecular studies, it is not possible to tell which one was which. Therefore, these must remain species inquirendae until they can be differentiated and named. There are at least three Sarcocystis species that have been named from gazelles, but only from sarcocysts in their muscle tissues. All three, S. gazellae Balfour, 1913 , S. mongolica Matschoulsky, 1947a ,b, and S. woodhousei Dogel, 1916 , are mentioned in Levine (1986 ) and in Dubey et al. (2015a , but the carnivore definitive host is not known for any of them. Clearly a lot of work still needs to be done just sorting out the various Sarcocystis species in gazelles. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Lucio-Forster and Bowman (2011) surveyed fecal samples of cats from two shelters (1,272 samples) and their affiliated foster homes (50 samples) in upstate New York, USA. Sarcocystis sporocysts were detected in 11/1,322 (<1%) samples. Determination was based on microscopy and morphology, and there was no attempt to identify to species. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. McKenna and Charleston (1980a) working on the North Island, New Zealand, examined fecal samples from domestic cats and identified sporocysts of Sarcocystis in 86/508 (17%) samples. There was no attempt to identify to species. Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Munday et al. (1978) examined muscle samples from 1,497 Australian mammals including 3 red foxes (Vulpes vulpes) and 27 brown fur seals (Arctocephalus pusillus (syn. S. doriferus)), but all 30 were negative. They also examined intestinal mucosal scrapings or feces from 55 feral cats and found small numbers of sporocysts in 1, and these measured 13-14 × 8.5. They (1978) concluded the sporocysts were "typical of Sarcocystis or Frenkelia." Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Ogassawara et al. (1986) examined fecal samples of domestic cats from different areas of São Paulo, Brazil and reported finding sporocysts of Sarcocystis in 18/215 (8%) fecal samples, mostly in animals 4-6-months-old (their Table 2 ). Definitive host: Felis catus L., 1758, Domestic Cat. Intermediate host: Unknown. Remarks: Serra et al. (2003) examined fecal samples of 65 household and 66 stray cats in Rio de Janeiro, Brazil. Sarcocystis sporocysts were seen in 1/131 (<1%) cats, from one household. Original hosts: Leopardus pardalis (L., 1758), Ocelot; Leopardus tigrinus (Schreber, 1775), Oncilla or Little Spotted Cat; Leopardus wiedii (Schinz, 1821) Margay. Remarks: Metzger et al. (2008) collected blood samples from 29 non-domestic neotropical felids in northeastern Brazil and found an ocelot, a little spotted cat, and a margay infected with an Hepatozoon species. They confirmed each infection by light microscopy and molecular analysis of partial sequences of the 18S rRNA gene of Hepatozoon, but no specific identification was attempted. Original host: Leopardus pardalis (L., 1758), Ocelot. Remarks: Patton et al. (1986) did a coprological survey of parasites of wild neotropical Felidae from the Cockscomb Basin, Belize, Central America and said they found oocysts of an Isospora species in 3/8 (37.5%) fecal samples from ocelots. No other information was given. Original host: Leptailurus serval (Schreber, 1776) (syn. Felis serval Schreber, 1776), Serval. Remarks: Mackinnon and Dibb (1938) looked at the feces of several serval at the London Zoo and reported oocysts they identified as those of I. felis. However, the oocysts they measured were 26-33 × 22-27, with sporocysts only 13 in mean length. They indicated the oocysts were partially sporulated when passed by their hosts. However, both the oocysts, and especially the sporocysts, are too small to be those of I. felis (now C. felis) and they are larger than those of I. rivolta (now C. rivolta). Levine and Ivens (1981) suggested these oocysts belong to a separate (unknown) species. We agree. It can only be relegated to a species inquirenda. Remarks: Bjork et al. (2000) published, "the first documentation of enteric parasites in a wild population of African lions in the Serengeti region" of northern Tanzania, Africa. They collected 33 freshly deposited fecal samples from wild lions, preserved them in 10% formalin, and examined both direct smears and fecal flotations for transmission stages. They said they found the following apicomplexans: Eimeria sp. in 1 lion, Isospora felis in 16, I. rivolta in 2, Isospora sp. in 1, Sarcocystis sp. in 15, and Toxoplasma-like oocysts in 4 samples. Unfortunately, they made these spurious identifications by measuring only one or a few oocysts, and all oocysts (except the oocysts/sporocysts of Sarcocystis) were unsporulated, which, of course, makes their identifications to genus completely unreliable, if not impossible. Original hosts: Panthera pardus (L., 1758), Leopard; Panthera tigris (L., 1758), Tiger; Panthera tigris bengalensis (L., 1758), Bengal Tiger. Remarks: Patton and Rabinowitz (1994) did a coprological survey of wild felids in Thailand and preserved the feces in 10% formalin, but made no mention that they took time to sporulate potential oocysts that may have been in their samples. Nonetheless, they reported finding Isospora-like oocysts that measured 40 × 32 in 2/54 (4%) P. pardus and in 1/3 (33%) P. t. bengalensis; Isospora-like oocysts that were 20 × 20 in 4/54 (7%) P. pardus; Toxoplasma-like oocysts in 1/54 (2%) P. pardus; Sarcocystis spp. oocysts/ sporocysts in 11/54 (20%) P. pardus and 7/19 (37%) P. tigris. These generalist determinations to genus seem completely unreliable to us. Original host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion. Remarks: Müller-Graf (1995) conducted a coprological survey of intestinal parasites of wild P. leo in the Serengeti and the Ngorongoro Crater, Tanzania, East Africa. She examined feces from lions and found "Coccidia" in 59/112 (53%) samples. No other information was given on the oocysts she saw. Original host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion. Remarks: Dubey and Jardine (2008) examined formalin-fixed tissues of a <2-day-old captive-born lion cub, P. leo, from Pretoria, South Africa. Meronts, merozoites, gamonts, and unsporulated oocysts were found in epithelial cells of the cub's ileum and examined by LM and TEM. Endogenous stages did not stain with antibodies to T. gondii and/or N. caninum and the morphology and size of various tissue stages eliminated I. (syn. C.) felis and I. (syn. C.) rivolta as suspects. Stages also differed from those of H. hammondi, and H. hammondi is not known to be transmitted transplacentally, nor are felids suspected to be definitive hosts of N. caninum. The age of the cub and the advanced development of the parasite supported the assumption that it was acquired in utero but no identification was/could be made. This seems a great opportunity for a future research project but, until then, it must be considered a species inquirenda. Original host: Panthera pardus (L., 1758), Leopard. Remarks: Karanis et al. (2007) examined feces of P. pardus in the Xining Zoo, Qinghai province, China, using an immunofluorescence test, nested PCR, and sequencing of a partial 18S rRNA gene; they reported Cryptosporidium oocysts in P. pardus and said they were C. parvum mouse genotype. Neither the number of examined leopards nor their origin was given. Original host: Panthera tigris sumatrae, Sumatran Tiger. Remarks: Lim et al. (2008) said they found oocysts of a Cryptosporidium sp. in the feces of 1/3 (33%) Sumatran tigers examined at Zoo Negra, Kuala Lumpur, Malaysia. No other information was given. Original host: Panthera (?), identified only as "leopard." Remarks: Wang and Liew (1990) reported oocysts of Cryptosporidium in a "leopard" from a bird park in Taiwan; the oocysts were detected in a fecal smear stained with modified Ziehl-Neelsen karbolfuchsin. No other information was given. Original host: Panthera tigris (L., 1758), Tiger (?). Remarks: Rastégaïeff (1929a) in a "note preliminaire" reported finding oocysts of this form in the feces of a tiger (species name not given) of the Zoological Gardens of Leningrad. She described the oocysts as spheroidal, 14-15 wide, without a M. She gave no other structural information other than it had four sporocysts. Rajasekariah et al. (1971) reported that they found this same "species" in a captured "panther" cub kept at the Dharmaram College, Bangalore, India and that it had spheroidal oocysts, 18-20 wide, with a 2-layered wall, ∼2.0 thick, and without a M, OR, PG, SB, SR but that it had ellipsoidal SP, 10 × 6, with SZ that measured 8 × 4. They also reported another eimerian, E. anekalensis, which they described as new, also from this panther (see Chapter 10, Felidae). It is questionable whether this poorly described form can be a parasite of both the tiger and the leopard. It is likely not a parasite of either, in our opinion. four samples from zoo lions, two of which had isosporan oocysts in them. Ovoidal oocysts were L × W (n = 30): 26 × 22 (23-33 × 20-28), L/W ratio: 1.2 (1.1-1.4); the oocyst wall was 2-layered, ∼1.3 thick, and lacked M, OR, PG. Sporocysts were ellipsoidal, 17 × 12 (16-18 × 11-13), L/W ratio: 1.4; SB, SSB, PSB: all absent, but SR was present and SZ were banana-shaped, 13 × 4 (12-15 × 3-9), with a central N and a prominent RB at the broad end. Pande et al. (1970) compared their sporulated oocysts to those of I. felis, I. laidlawi, and I. leonine and thought theirs was sufficiently different. Nonetheless, they wrote, "It is not possible to assign present material categorically to any of these" and, "pending the availability of more material, these oocysts are, therefore described under an unnamed species, Isospora sp." Levine and Ivens (1981) questioned whether or not this is a true parasite of the lion, and so do we. This morphotype has not been seen or described since its original discovery, but if it is found again and, indeed, is a true parasite of the lion, it must be placed in the Cystoisospora. with numerous crescentic bodies" in sections of heart muscle from two zoo lions (presumably P. leo) that had died in India. The sarcocysts, examined only under LM, were, 12.3 × 2.4, did not have "discernable" septa, and they were found in both striated and cardiac muscle fibers and in the cells of Purkinje fibers. Because of the small size and lack of septa (?), it is unlikely that this form can be attributed to S. felis, so it must be considered a species inquirenda. Bhatvdekar and Purohit (1963) said they fed sarcocysts in muscle tissue to mice and injected "the spores" (bradyzoites?) intramuscularly into mice and that "typical sarcocysts developed within four to seven weeks after feeding the infective stages." This remains to be confirmed. Definitive host: Unknown. Intermediate host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion. Remarks: Bwangamoi et al. (1990) found numerous sarcocysts measuring 730-760 × 113-114 in the skeletal muscles of a 7-year-old lioness from Nairobi National Park, Africa. No other information was given, thus, a species inquirenda. Definitive host: Panthera tigris (L., 1758), Tiger. Intermediate host: Unknown. Remarks: Jog et al. (2003) surveyed the feces of 69 tigers from Tadoba National Park, Maharashtra, India, and found 3/69 (4%) passed sporocysts of a Sarcocystis species, and 2/36 (5.5%) tigers from Mudumalai National Park and Wildlife Sanctuary passed similar sporocysts in their feces. No attempt was made to further characterize the species, but Jog et al. (2003) also mentioned that 0/28 leopards, presumably Panthera pardus (L., 1758), were never found to have sporocysts in their feces when examined. Definitive host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion. Intermediate host: Unknown. Remarks: Müller-Graf (1995) did a coprological survey of wild P. leo in the Serengeti and the Ngorongoro Crater, Tanzania, East Africa. She examined feces from lions and found "Coccidia" in 59/112 (53%) samples. She also reported finding sporocysts representing a Sarcocystis species in 16/112 (28%) samples. No other information was given on these samples. Definitive host: Panthera pardus (L., 1758), Leopard. Intermediate host: Unknown. Remarks: Patton and Rabinowitz (1994) did a coprological survey of wild felids in Thailand and found Sarcocystis sporocysts in the feces of 11/54 (20%) samples examined. No other information was presented; thus, another species inquirenda. Definitive host: Panthera tigris (L., 1758), Tiger. Intermediate host: Unknown. Remarks: Patton and Rabinowitz (1994) did a coprological survey of wild felids in Thailand and found Sarcocystis sporocysts in the feces of 7/19 (37%) samples examined. No other information was presented, thus another species inquirenda. Definitive host: Unknown. Intermediate host: Panthera pardus (L., 1758), Leopard. Remarks: Pellérdy (1974b) proposed this name for I. rivolta from the cat because he believed I. rivolta was first discovered and named from the dog. However, Wenyon (1923) reviewed all of the literature on the coccidia known to that date from dogs and cats and convinced most colleagues that I. rivolta was first discovered in the cat. Thus, as concluded by Dubey (1975c), and we agree, this renders I. novocati to be rendered a nomen nudum. Original host: Martes martes (L., 1758), European Pine Marten. Remarks: Galli-Valerio (1932) said he saw an ovoidal coccidium that was, 7 × 2.25 with a flattened micropyle in the feces of the European pine marten and that it developed two sporocysts each with four sporozoites. We know of no oocysts in any apicomplexan family that has oocysts this small. No other structural data were given nor did he include a line drawing. Therefore, this name must be considered a nomen nudum. novocati of Pellérdy, 1974b, and Isospora mustelae of Galli-Valerio, 1932 , are considered nomena nuda, by definition. If one takes time to look into any of the individual entries cited above in this chapter, you might imagine each as the beginning of a short story that may wet your appetite for more information, but the story ends and leaves you wanting to know more. Even though many of these citations may seem trivial on first examination, they all present some modest baseline data, and virtually any one of these efforts could be expanded on by the right person(s). From the above numbers, two things may be obvious; (1) most carnivore populations have been poorly surveyed for apicomplexan parasites; and (2) there are a plethora of apicomplexan genera and species to be discovered, studied in much more detail, and be defined in many different ways. We are just now beginning to have a basic understanding of most of the apicomplexan genera mentioned in this chapter and we now know where and how to look for them. For example, Besnoitia species are often overlooked or misidentified as sarcocysts of Sarcocystis when carcasses are examined. It is still worthwhile and useful to study the oocysts of Eimeria, Isospora, Caryospora, and Cystoisospora species, but they must be carefully defined by measurements, photomicrographs, and line drawings. Sporocysts of Isospora species must be examined carefully to determine the presence or absence of a Stieda body. If a Stieda body is present, crosstransmission and feeding experiments will be needed to determine if sporulated oocysts can infect the host species in which they were discovered, or their nearest relatives, or the original oocysts may belong to a prey item and simply be passing through the gut of the original host from environmental contamination. If the sporocysts of larger isosporan-like oocysts do not have a Stieda body, it is most likely they are a species of Cystoisospora and lab mice should be infected to determine if they can act as intermediate/transport hosts for some asexual stage(s). Lots of surveys look at certain muscles of potential host species for the presence of sarcocysts and if there are none present (or none are "seen") in the specific muscles examined, the animal necropsied is considered negative for Sarcocystis. However, those doing surveys should understand that some sarcocysts are not visible to the naked eye, especially if they are very young, some are very small even when mature and can best be found by pepsin digestion versus gross inspection, and many species have their highest densities in different tissues; to some, this may seem reminiscent of how Eimeria species seem to divide up the intestinal tract, each destined to live in its own site-specific location. To illustrate the point we are making, Mohammed (2000) and Mohammed et al. (2000) looked for Sarcocystis infections in different gazelle species at the King Khalid Wildlife Research Centre in Saudi Arabia. They detected no macroscopic sarcocysts by fibre optic examination, a 40% infection rate was detected by histological examination, and an overall prevalence of infection of 67% was reached when they used pepsin digestion of muscle. In addition, the esophagus, diaphragm, heart, tongue, and skeletal muscles had different rates of infection dependent on the host gazelle species that was examined. They also reported that Sarcocystis infection was significantly higher in free-ranging gazelles kept in a main enclosure than in gazelles kept in breeding pens and higher in adult versus juvenile gazelles. Thus, there are lots of ways to examine every problem and, with the plethora of inquiries presented in this chapter, there are lots of problems awaiting future study. illustrated only a gamont/gametocyte in a granulocyte (PMN) of an east African lion; its size was given as 8.5 (8-10). DISCUSSION In the Caniformia, there are no casual, common-name-only, or genus-name-only references in these family and genera: Canidae: Atelocynus Cabrera 1857 (2 species); Mephitidae: Conepatus Gray, 1837 (4 species) Mustelidae: Aonyx Lesson 1827 (2 species) Hydrictis Pocock Galictis Bell, 1826 (2 species) species) Poecilogale Thomas, 1883 (monotypic) Odobenidae: Odobenus Brisson, 1762 (monotypic); Otariidae: Arctocephalus É. Geoffroy Saint-Hilaire, and Phocarctos Peters, 1866 (monotypic); Phocidae: Halichoerus Nilsson, 1820 (monotypic) 1844 (monotypic); Procyonidae species) species) 1915 (monotypic); and Ursidae: Melursus Meyer In total, there is no record that 37/72 (51%) Caniformia genera, and their 60 species (above), have any abbreviated coccidian identifications associated to them. And, in the 35 genera that have been surveyed 5%) species in those genera have been looked at sufficiently to have various coccidian forms known from them. In the Feliformia, we found no casual, common-name-only, or genus-name-only references in these family and genera: Eupleridae: Cryptoprocta Bennett 1865 (monotypic); Felidae: Caracal Gray, 1843 (monotypic) species) Uncia Gray, 1854 (monotypic) Hyaenidae: Hyaena Brisson, 1762 (2 species) Geoffroy Saint-Hilaire, 1824 (monotypic); Nandiniidae: Nandinia Gray, 1843 (monotypic); Viverridae: Arctictis Temminck 1897 (monotypic) Diplogale Thomas, 1912 (monotypic) species); and Viverricula Hodgson, 1838 (monotypic) have any abbreviated coccidian identifications associated to them. And, in the 17 genera that have been surveyed, only 27/71 (38%) species in those genera have been looked at sufficiently to have various coccidian forms known from them. In total, we found 483 apicomplexans that have been mentioned in the literature and placed into 17 genera or generic categories, but that either have not been described sufficiently or there was so little information provided by the author(s), that their validity, and sometimes even their identity was in question 34 Coccidia-like or Coccidia spp., 135 Cryptosporidium spp., 2 Cyclospora spp 36 Eimeria spp., 3 Hammondialike forms; 7 Hammondia-Neospora-like forms Hammondia-Toxoplasma-like forms; 14 Hepatozoon spp., 1 Hoareosporidium sp., 72 Isospora spp 135 Sarcocystis spp., and 3 Toxoplasma-like forms. All are considered species inquirendae Definitive host: Unknown. Intermediate host: Pusa caspica (Gmelin, 1788) (syn. Phoca caspica), Caspian Seal. Remarks: Kuiken et al. (2006) , reported on a die-off of Caspian seals in the spring and summer, 2000, which they attributed to canine distemper, and mentioned finding Sarcocystis-like cysts containing many banana-shaped bradyzoites, ∼6 μm × 2 μm, with a distinct N, in myocytes of the esophageal muscularis and skeletal muscle of 1/18 (5.5%) seals. The cysts distended the infected myocyte but did not induce an inflammatory response in adjacent tissue (their Fig. 12, p. 328) . No other information was provided. This species has not been identified since 2006 and must remain a species inquirenda. Definitive host: Unknown. Intermediate host: Pusa hispida (Schreber, 1775), Ringed Seal. Remarks: Migaki and Albert (1980) found sarcocysts in the skeletal muscles, but not in the heart, of 12/18 (67%) apparently healthy P. hispida from the Arctic Ocean near Barrow, Alaska, USA. The cysts were compartmented, elongate, 60-550 wide and had moderately thick, welldefined cell walls, 0.8-1.0 thick. Bradyzoites were 10-12 × 2-3. Obviously, the life cycle of this species is unknown. No other information was provided. This species has not been identified since 1980 and must remain a species inquirenda. Definitive host: Unknown. Intermediate host: Potos sp., Kinkajou. Remarks: Takos (1957) found a sarcosporidian infection in an old male kinkajou shot near the Rio Mandinga Bridge, Panama Canal Zone, Panama. Sarcocysts were found in striated muscle, but not in the myocardium or other organs. Sarcocysts were broadly ovoidal and measured 123-216 × 85-100. Each sarcocyst was bounded by a dense, well-defined, unstriated wall, ∼1 thick. Sarcocysts were divided into compartments by thin septa and the chambers they formed were densely packed with bananashaped zoites that measured 7.5-9 × 2. Takos (1957) said there was no evidence of tissue reaction to the parasite and that this was the first report of a Sarcocystis species in the muscles of a kinkajou. No other information was given. Original host: Procyon lotor (L., 1758), Raccoon. Remarks: Carlson and Nielsen (1982) reported the presence of "macrogametocytes containing multiple substructures identified as macrogametes and red-staining residual bodies, as well as oocysts" in the villi and intestinal lumen of the Definitive host: Felis catus L., 1758, Domestic Cat (?).Intermediate host: Mus musculus L., 1758, House Mouse. Remarks: Wallace (1975) looked at two morphologically different cysts in mouse skeletal muscles, which resulted when he inoculated them "with fecal material from a stray cat containing Isospora-type oocysts." The most common cyst type turned out to belong to T. gondii, which he confirmed by feeding the cysts to lab cats that produced oocysts that were 13 × 11 and resembled those of T. gondii. The second type of cyst found in mouse skeletal muscles was "observed in only a few mice, contained bradyzoites resembling those of Sarcocystis, but the oocyst or sporocyst that gave rise to it was overlooked and apparently lost." Original host: Felis catus L., 1758, Domestic Cat. Remarks: Epe et al. (2004) , in northern Germany, surveyed fecal samples of cats from 1998 to 2002 and found 3/441 (<1%) to pass Toxoplasma-like oocysts, which did not "fit" in any particular genus. Original host: Felis catus L., 1758, Domestic Cat. Remarks: Christie et al. (1976) surveyed intestinal parasites of cats, all > 3-months-old, from a shelter in Columbus, Ohio, USA. "Toxoplasmalike" oocysts, 12 × 11, were detected in 9/1,000 (0.9%) cats. The oocysts were orally inoculated into mice, then the inoculated mice were fed to SPF cats; 1 SPF cat began to shed similar oocysts in its feces. Original host: Lynx rufus (Schreber, 1777), Bobcat.Remarks: Carver et al. (2012) collected fecal samples from 141 bobcats in California (Ventura county) and Colorado (Front Range and Western Slope), USA. Samples were examined by flotation and direct immunofluorescence followed by PCR assays, but the authors did not specify what genes were amplified (!) and the PCR amplification failed. The authors said they detected Cryptosporidium across all studied sites but did not provide the number of positive animals nor the prevalence. Original host: Lynx rufus (Schreber, 1777), Bobcat.Remarks: Ziegler et al. (2007) examined fecal samples from bobcats in southeastern New York state, USA, by flotation and LM, and by polyclonal Cryptosporidium antigen-capture ELISA (considered positive based on an optical density ≥0.050). Cryptosporidium was found in 1/14 (7%) samples, but not identified to species. Original host: Lynx lynx isabellinus (Blyth, 1847), Central Asian Lynx. Remarks: Levine and Ivens (1981) listed this eimerian as having been described by Anpilogova and Sokov (1973) from Tadzhikistan (former USSR). They said the ellipsoidal oocysts measured 33 × 23 (30-38 × 19-27) and had a bilayered wall with a M in the inner wall. However, neither pair of authors provided a line drawing or photomicrograph; thus, it can only be a species inquirenda. Original host: Lynx lynx isabellinus (Blyth, 1847), Central Asian Lynx.Remarks: Levine and Ivens (1981) listed this eimerian as having been described by Anpilogova and Sokov (1973) from Tadzhikistan (former USSR). They said the elongate-ovoidal oocysts were truncated at the M and measured 40.5 × 27, with sporocysts that were 13.5 × 11, with no other information. Neither pair of authors provided a line drawing or a photomicrograph; thus, it can only be considered a species inquirenda. Original host: Lynx lynx isabellinus (Blyth, 1847), Central Asian Lynx.Remarks: Levine and Ivens (1981) listed this eimerian as having been described by Anpilogova and Sokov (1973) from Tadzhikistan (former USSR). They said the ovoidal oocysts measured L × W: 31.0 × 23.5 (24-32 × 19-27), with a 2-layered wall and without a M, OR, PG. Neither pair of authors provided a line drawing or photomicrograph; thus, a species inquirenda. Original host: Lynx rufus (Schreber, 1777), Bobcat. (1983) (1976) found oocysts they called I. bigemina large form and were among the first to use them to look at in vitro excystation of sporocysts that lack Stieda bodies. The oocysts and sporocysts were, of course, an unknown species of Sarcocystis. Although the species from the bobcat remains a species inquirenda, the interesting observation was that during excystation, the walls of the sporocyst collapsed, apparently along predetermined lines, allowing sporozoites to escape. As the sporocyst collapsed, the wall separated into two halves, each with two pieces attached together at a point corresponding to one pole of the original sporocyst. Synonym: Isospora felis (Wasielewski, 1904) Wenyon, 1923.Original host: Lynx sp., Bobcat.Remarks: Triffitt (1927) said she found oocysts of an Isospora in the rectal contents of a lynx that died in the Zoological Gardens, London, but did not specify the host species (L. lynx or L. canadensis). She did state that its oocysts were identical to those of I. felis, but that they had a M, which those of I. felis lack. Based on that difference, Levine and Ivens (1981) named the form partially described by Triffitt (1927) as I. lyncis because she also gave some mensural data. Triffitt (1927) said the oocysts were ovoidal, 40-47 × 28-37, and had an oocyst wall with 3-layers, ∼0.75 thick, and a M, ∼4-5 wide; ovoidal SP were 20-33 × 14-18, with a SR, and SZ measured 15 long, with a large RB at their more rounded end and a smaller one at their more pointed end. Triffitt (1927) said that sporulation was completed in 7-9 days at room temperature. However, neither Triffitt (1927) nor Levine and Ivens (1981) provided a line drawing or a photomicrograph. Thus, there is no evidence that this species actually exists, so the name only can be a species inquirenda. Original host: Puma yagouaroundi (É. Geoffroy Saint-Hilaire, 1803), Jaguarundi. Remarks: Metzger et al. (2008) collected blood samples from 29 non-domestic neotropical felids in northeastern Brazil and found a jaguarundi infected with an Hepatozoon species. They confirmed this infection by light microscopy and molecular analysis of partial sequences of the 18S rRNA gene of Hepatozoon, but no specific identification was attempted. Definitive host: Puma (syn. Felis) concolor (L., 1771), Puma, Cougar.Intermediate host: Unknown.Remarks: Dubey (1982b) surveyed feces for coccidians from a variety of wild carnivores in Montana, USA and reported sporocysts of Sarcocystis in 2/12 (17%) pumas. Sporocysts were 11.5-13.0 × 7.5-8.8. No attempt was made to identify the species beyond genus; thus, a species inquirenda. Definitive host: Unknown. Intermediate host: Puma (syn. Felis) concolor (L., 1771), Puma, Cougar. Remarks: Kluge (1967) found sarcocysts in the skeletal muscles of a 13-year-old puma that died in the National Zoological Park, Washington, D.C., USA. Sarcocysts, examined with LM, had a thin wall, composed of a single layer without septa projecting from the internal surface. The cyst contained numerous banana-shaped structures (bradyzoites?), "each of which had a round to oval subterminal nucleus and small basophilic granules within the cytoplasm." Kluge (1967) said these cysts "replaced approximately two-thirds of the sarcoplasm in the slightly swollen affected muscle bundles," but that no inflammatory reaction was seen. No other information and no measurements were given, but because he reported "no septa" to be present, it is unlikely that this form can be attributed to S. felis, so it must be considered a species inquirenda. GENUS PANTHERA (L., 1758) (4 SPECIES) Original host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion. AND BROCKLESBY, 1971Original host: Panthera pardus (L., 1758), Leopard.Remarks: Keymer and Brocklesby (1971) described intraleukocytic gametocytes and extracellular forms of a Hepatozoon species in the blood, and two forms of meronts of a Hepatozoon in P. pardus from central Africa. Meronts were found, both in cardiac muscle and lungs. They said that the gametocytes were indistinguishable from H. canis of the dog, but unlike the Hepatozoon of the dog, no meronts were found in liver, bone marrow, spleen, or lymph nodes. This was the first description of Hepatozoon meronts from a leopard, but its actual identity remains a mystery. Original host: Panthera leo massaica (Neumann, 1900), Massai Lion. Remarks: Krampitz et al. (1968) said that about 28/56 (50%) Massai lions had "gametocytes in its monocytes, which differed from all other species described so far." Original host: Panthera pardus (L., 1758), Leopard. Remarks: Mandal and Choudhury (1983) collected feces from one P. pardus, in the Betla Forest Palamau Tiger Reserve, Bihar, India. They incubated the sample in 2%-2.5% potassium dichromate solution and concentrated oocysts via centrifugation. They found ovoidal isosporan oocysts with a 2-layered wall, ∼0.9 thick, that measured, L × W: 29.2 × 23.4 (26-31 × 22-24), L/W ratio: 1.2; M, OR, PG: all absent. Sporocysts were spheroidal, 15.3 (12-17); L/W ratio: 1.0; SB, SSB, PSB: all absent, but SR was present as a large, granular, spheroidal mass, 5.8 (5-7) wide; SZ were banana-shaped, 10.3 × 2.6 (7.5-10 × 2-3), with a prominent RB at the broad end. They chose only to identify the oocysts measured as I. rivolta-like. Its true identity is anyone's guess. Original hosts: Panthera leo (L., 1758), Lion; Panthera tigris (L., 1758), Tiger.Remarks: Chauhan et al. (1973) did a "preliminary survey" of parasitic infections among mammals and birds at the Lucknow Zoological Garden, India and reported finding Isospora sp. oocysts in the feces of P. leo and P. tigris, but the number of lions and tigers examined was not stated. Original host: Panthera tigris altaica Temminck, 1844, Tiger. Remarks: Hou et al. (2008) found coccidian oocysts in a naturally-infected tiger in China; based on morphology, sporulation, and phylogenetic analysis, they concluded the coccidian from their tiger was Isospora. No other information was given.Original host: Panthera pardus (L., 1758), Leopard. Remarks: Patton et al. (1986) did a coprological survey of neotropical felids in Belize; they found oocysts of an Isospora species in the feces of 2/25 (8%) jaguars from the Cockscomb Basin of Belize, Central America. No other information was presented. Original host: Panthera pardus (L., 1758), Leopard.Remarks Definitive host: Unknown.Intermediate host: Panthera leo (L., 1758) (syn. Leo leo Frisch, 1775), Lion.Remarks: Bhatavdekar and Purohit (1963) may have been the first to document sarcocysts in the muscles of felids. They said they found, "a very thin hyaline capsule and the cavity was filled Remarks: Somvanshi et al. (1987) examined the carcass of a young leopard that was shot in Ramgarh, Uttarakhand, India, in the foothills of the Himalayas. Numerous elongated sarcocysts were found in the myocardium; the sarcocyst wall was thin, with numerous trophozoites. In spite of the large number of sarcocysts seen, the authors reported no inflammatory reaction. No other information was presented; thus, another species inquirenda. Original host: Cynictis penicillata (G.[Baron] Cuvier, 1829), Yellow Mongoose. Remarks: Markus (1972) reported isosporanlike oocysts from the yellow mongoose in South Africa, but no measurements were given. Definitive host: Unknown.Intermediate host: Helogale parvula (Sundevall, 1847), Common Dwarf Mongoose.Remarks: According to Levine and Ivens (1981) , Viljoen (1921) found sarcocysts in the striated muscles of H. parvula in South Africa, but they did not see that actual paper, and got the citation from Nietz (1965). Levine (1986) did not list this species, and Dubey et al. (2015a) listed neither the Viljoen nor Nietz references and made no mention of a Sarcocystis species in H. parvula. Viljoen (1921) does not come up in Google Scholar and we were unable to obtain a copy of the Nietz paper. Original host: Herpestes ichneumon (L., 1758), Egyptian Mongoose. Remarks: Balozet (1933) found oocysts in fecal matter of two H. ichneumon from the Zaghousan region, Tunisia. He measured 100 sporulated oocysts and offered a very superficial description saying the subspheroidal oocysts were L × W: 22 × 19 (19-26 × 16-20) , with a transparent, 2-layered wall and a poorly visible M. He said oocysts sporulated in 4 days when fecal material was placed on filter paper impregnated with potassium dichromate. Balozet (1933) fed sporulated oocysts in fecal material to a 4-dayold puppy and 9 days later, reinfected it with more oocysts. Four days after the second inoculation, oocysts were "extremely numerous" in the dog's feces, and the pup was described as having dysentery; after the fourth day of patency, oocysts became rare in the feces and the pup was sacrificed 30 days after the first inoculation. Balozet (1933) found no visible intestinal lesions and concluded that he had found Isospora rivolta (Grassi, 1879) Wenyon, 1923, saying, "…which appears to be a rather ubiquitous species capable of parasitizing several species of carnivorous species." In spite of this very modest description, and the lack of line drawing or photomicrograph, Levine et al. (1975) Remarks: According to Levine and Ivens (1981), Viljoen (1921) found sarcocysts in the striated muscles of M. mungo in South Africa, but they did not see Viljoen's paper and got the citation from Nietz (1965) . Levine (1986) Original host: Crocuta crocuta (Erxleben, 1777), Spotted Hyaena. Remarks: Krampitz et al. (1968) said that about 4/9 (44%) spotted hyaenas in the Serengeti National Park, Tanzania had gametocytes of a Hepatozoon species in their monocytes and neutrophiles. Original host: Crocuta crocuta (Erxleben, 1777), Spotted Hyaena. Remarks: Berentsen et al. (2012) surveyed hyaenas and other African carnivore species from the Luangwa Valley, Zambia for gastrointestinal parasites; they reported Isospora species in 3/9 (33%) C. crocuta but gave no other information about this parasite. They collected fecal samples of 108 civets from December, 2014 to March, 2015; samples were taken from 10 "forest fragments" that ranged from 8-2,000 ha. All samples were collected in 10% formalin, and they (2016) could not identify to which of the three endemic civets each fecal sample belonged. Samples thought to be infected with a Cyclospora sp. were found in two forest fragments, those infected with Eimeria species were found in four forest fragments and those identified only as Coccidia species were found in seven forest fragments. It is not clear how they defined "Coccidia" species nor how they were able to distinguish oocysts of Cyclospora and Eimeria species, given that feces were collected in 10% formalin and, apparently, not placed into thin layers in Petri dishes to allow sporulation. Original host: Paradoxurus hermaphroditus (Pallas, 1777), Asian Palm Civet.Remarks: Colón and Patton (2012) did a coprological survey of civets in Sabah, Borneo, and feces from a road-killed P. hermaphroditus was reported to have oocysts of an Eimeria species in its feces. No other information was given. Original host: Genetta genetta (L., 1758), Common Genet. Remarks: Mateo et al. (2017) examined fecal samples of genets, from three autonomous regions of Spain (Basque Country, Castile-La Mancha, Extremadura) and examined them by nested PCR targeting the 18S rRNA gene, followed by genotyping based on the gp60 gene. Cryptosporidium was detected in 1/6 (17%) samples, the one from Extremadura. Original host: Genetta dongolana (L., 1758) (syn. Genetta dongolana Hemprich and Ehrenberg, 1832), Common Genet. Remarks: Agostinucci and Bronzini (1953) examined the intestinal contents of one G. dongolana that died in the Zoological Garden of Rome, Italy, a few days after it arrived from Somalia (East coast of Africa) and found unsporulated oocysts. They kept the fecal material in 3% potassium dichromate solution at ∼18°C and said, "the oocysts reached full maturity in ∼7-8 days." Sporulated oocysts were slightly ellipsoidal and "featured a micropyle in each of the poles, with rather thin-walled, smooth, and seemingly double outline." The oocysts were L × W: 25.4 × 19.9 (20-30 × 12.5-25) , L/W ratio: 1.3; OR, PG: both absent. Sporocysts were reported to be "pear-shaped pods," 8.4 × 6.0 (6-12.5 × 5-7.5); SR: absent. There was no mention of presence or absence of SB, SSB, PSB, and there was no line drawing or photomicrograph, presented as a type, to support this inadequate description. No one in the literature has referred to this species since its original description. Therefore, it must be relegated to a species inquirenda. Original hosts: Genetta genet (L., 1758), Common Genet; Genetta thierryi Matschie, 1902 (syn. Genetta rubiginosa Pucheran, 1855), Haussa Genet; Genetta tigrine (Schreber, 1776), Cape Genet.Remarks: Keymer and Brocklesby (1971) studied and described intraleukocytic and extracellular stages of a Hepatozoon sp. they found in the blood of G. rubiginosa and G. tigrina, and meronts in the liver and cardiac muscle of G. rubiginosa. They found two forms of meronts in the heart, and both differed from meronts found in the liver. This was the first time Hepatozoon meronts have been described in Genetta spp. Original host: Viverra tangalunga (Gray, 1832) , Malayan Civet. Remarks: Colón and Patton (2012) did a coprological survey of civets in Sabah, Borneo, and a female V. tangalunga was reported to have Eimeria and Isospora oocysts and sporocysts of a Sarcocystis sp. in her feces. But no other information was given. Some authors, especially in some of the very old literature, gave new names to organisms they saw, but for which they presented no other substantive information. Such a name is called a "nude name" or nomen nudum (pl. nomena nuda). These names become preoccupied and unavailable names; however, the same name may be made available at a later time for the same or a different concept, but in such case, it would take authorship and date from the act of establishment, not from the earlier publication as a nomen nudum. Original host: Felis catus L., 1758, Domestic Cat.