Compiled by
Donald W. Duszynski¹, Lee Couch¹, and Steve J. Upton^2
1Department of Biology, University of New Mexico
²Division of Biology, Kansas State University

NOTE: The coccidia of canids and felids represent an exceptionally confused group of coccidia. Some species are morphologically distinguishable, early experimental transmission studies were clouded by poorly controled studies lacking coccidia-free animals, a number of species were poorly described and many are most likely synonyms, a wide number of misidentifications have cluttered the literature, some microscopes appear to have been miscalibrated, and a number of species have now been placed in other genera as they have been shown to utilize heteroxenous life-cycles. In addition, recent molecular evidence (Carreno and Barta 1999; Carreno et al. 1998) suggests that the genus Isospora is polyphyletic and that the isosporans of carnivores, primates, and even Isospora suis from swine are more closely related to members of the Sarcocystidae Poche, 1913 than to eimerians or other isosporans in the family Eimeriidae Minchin, 1903. Once additional species are examined, it seems likely that most isosporans of carnivores and primates will be moved out of the Eimeriidae and into the Sarcocystidae. Since the genus Cystoisospora Frenkel, 1977 has already been established for these coccidia, no new generic names should need to be created.

I. Named and potentially valid canid Isospora spp.

Synopsis: We consider only 4 valid Isospora spp. in domestic dogs. Isospora canis Nemeseri, 1959, the largest of the canine isosporans, is very common and quite distinctive. However, this species may in fact be the same as one or both of the morphologically similar species found in the fox and termed Isospora canivelocis (Weidman, 1915) Wenyon, 1923 and Isospora buriatica Yakimoff and Matschoulsky, 1940 in Matschoulsky, 1941. Either of these names would have priority over Isospora canis should cross-transmission studies reveal synonymy. The three species in the Isospora ohioensis-complex are also distinct species, although morphologically very similar. Isospora burrowsi Trayser and Todd, 1978, Isospora neorivolta Dubey and Mahrt, 1978, and Isospora ohioensis Dubey, 1975 are reported to have similar oocyst structure although the middle species has never had a proper description of the oocysts. It is plausible that Isospora pavlodarica Nukerbaeva and Svanbaev, 1973, most of the reports of Isospora vulpina Nieschulz and Bos, 1933 including the original description, and the report by Golemansky and Ridzhakov (1975) of Isospora vulpis Galli-Valerio, 1931 are actually the same as members of the Isospora ohioensis-complex. Here again, future cross transmission studies should resolve these problems. Unfortunately for traditionalists, the species reported from the domestic dogs would fall as synonyms to the older named species from the fox. There does appear to be one morphotype in the fox that falls between Isospora canis and the species in the Isospora ohioensis-complex. Bledsoe (1976a, 1976b) termed this species Isospora vulpina Nieschulz and Bos, 1933 whereas Galli-Valerio (1931) and Svanbaev and Rachmaatullina (1971) term this Isospora vulpis Galli-Valerio, 1931. The bothersome thing is that Bledsoe (1976a, 1976b) was able to readily infect domestic dogs. If this were a distinct species, and if Canis spp. can become readily infected, it is likely that these oddly sized oocysts would have been routinely noted in domestic dogs over the years. One possible explanation is that the sizes of the oocysts vary slightly in the different host species. There is precedence for this; for instance Eimeria dispersa Tyzzer, 1929 produces larger oocysts in quail than it does other galliforms.

1. Isospora buriatica Yakimoff and Matschoulsky, 1940 in Matschoulsky, 1941
   1. Hosts: Vulpes bengalensis (Indian fox), Vulpes corsac (Corsac fox - type host)
   2. Oocysts: 38.9 x 29.4 (26.4-41.3 x 23.1-31.4) micrometers. Svanbaev (1956) gives measurements of 25.8 x 19.0 and 31.7 x 24.8 micrometers to two different isosporans from the Corsac fox, the second of which comes the closest but is short of matching the original description. Dubey (1963) gives measurements of 31 x 25 (29-33 x 22-25), Nukerbaeva and Svanbaev (1973) 30.8-44.8 x 25.2-30.8, and Glebezdin (1978) 30.9 x 25.2 (30.8-32.2 x 25.2) micrometers.
   3. References: Dubey 1963; Matschoulsky 1941; Nukerbaeva and Svanbaev 1973; Svanbaev 1956; Yakimoff and Matschoulsky 1940
   4. Remarks: Various authors (Glebezdin 1978; Levine and Ivens 1981; Pellerdy 1974) reference Yakimoff and Matschoulsky (1940) as the original publication. However, no such name appears in the 1940 paper. Matschoulsky (1941) seems to be the first to report this coccidian and the confusion appears to lie with the fact that he credits the description as Yakimoff and Matschoulsky, 1940. Although perhaps a separate species, the original measurements provided by Matschoulsky (1941) suggest that this may be Isospora canis Nemeseri, 1959. If so, Isospora canis would be come a junior synonym. All other authors seem to have been working with oocysts representing smaller species and not Isospora buriatica. Svanbaev (1956) was obviously working with a mixed population of coccidia.

2. Isospora burrowsi Trayser and Todd, 1978
   1. Host: Canis familiaris (domestic dog - type host)
   2. Oocysts: 20 x 17 (17-22 x 16-19) micrometers
   3. References: Heine 1981; Rommel and Zielasko 1981; Trayser 1973; Trayser and Todd 1978
   4. Remarks: Oocysts of Isospora burrowsi tend to be smaller than those of Isospora ohioensis. The species reported by Arthur and Post (1977) from coyotes, Canis latrans, may actually represent Isospora burrowsi.

3. Isospora canis Nemeseri, 1959
   1. Synonym: Cystoisospora canis (Nemeseri, 1959) Frenkel, 1977
   2. Synonym: Diplospora bigemina Wasielewsky, 1904 pro parte
   3. Synonym: Levinea canis (Nemeseri, 1959) Dubey, 1977
   4. Host: Canis familiaris (domestic dog - type host); experimentally Canis latrans (Coyote); perhaps Vulpes vulpes (Red and silver foxes)
   5. Oocysts: 38 x 30 (34-42 x 27-33) micrometers. Nemeseri (1960) gave measurements as 39 x 32 (36-44 x 29-36), Levine and Ivens (1965) 38 x 30 (35-42 x 27-33), de Rocha and Lopes (1971) 36.3 x 30.0 (32.1-40.2 x 23.5-35.8), and Lepp and Todd (1974) 36 x 30 (34-40 x 28-32) micrometers. Oocysts from coyotes were measured as 36.8 x 30.7 micrometers by Loveless and Anderson (1975).
   6. References: Dubey 1973, 1982; Dunbar and Foreyt 1985; Heine 1981; Lepp and Todd 1974, 1976; Loveless and Anderson 1975; Markus 1983; Nemeseri 1959, 1960; Reinhardt 1934; de Rocha and Lopes 1971; Roberts et al. 1972; Stiles 1892; Streitel and Dubey 1976
   7. Remarks: Oocysts tend to be slightly egg-shaped (attenuated at one end) and are the largest and most distinctive of the isosporans in dogs. It seems possible that several of the reports of Isospora canivelocis (Weidman, 1915) Wenyon, 1923, including the original description, as well as Isospora buriatica Yakimoff and Matschoulsky, 1940 in Matschoulsky, 1941, may actually represent Isospora canis Nemeseri, 1959. In that case, Isospora canis Nemeseri, 1959 would fall as a junior synonym.

4. Isospora canivelocis (Weidman, 1915) Wenyon, 1923
   1. Synonym: Coccidium bigeminum var. canivelocis Weidman, 1915
   2. Synonym: Isospora bigeminum var. canivelocis (Weidman, 1915) Mesnil, 1916
   3. Synonym: Isospora canivecolis (Weidman, 1915) Wenyon, 1923, lapsus
   4. Synonym: Lucetina bigeminum var. canivelocis (Weidman, 1915) Henry and Leblois, 1926
   5. Hosts: Alopex lagopus (Arctic fox); Vulpes velox (Swift fox - type host); Vulpes vulpes (Red and silver foxes)
   6. Oocysts: 31-38 x 25-30 micrometers. Sprehn and Cramer (1931) measured oocysts as 21-38.4 x 16.8-31.2, Neischulz and Bos (1933) 36.1 x 29.7 (33-39 x 27-32), Svanbaev (1960) 27.4 x 26.1 (23.7-32.4 x 19.6-32.0), Nukerbaeva and Svanbaev (1973) 29.4 x 26.3 (28-30.8 x 25.2-28), Golemansky and Ridzhakov (1975) 35.0 x 27.0 (30.0-39.0 x 23.0-30.4), and Glebezdin (1978) 32.6 x 27.4 (28.0-36.4 x 25.2-30.8) micrometers.
   7. References: Glebezdin 1978; Golemansky and Ridzhakov 1975; Nieschulz and Bos 1933; Nukerbaeva and Svanbaev 1973; Ridala 1936; Sprehn and Cramer 1931; Svanbaev 1960; Weidman 1915
   8. Remarks: The original description by Weidman (1915) suggests that this coccidian may actually be Isospora canis Nemeseri, 1959. Measurements provided by Neischulz and Bos (1933) and Glebezdin (1978) suggest the same. The smaller sizes given by some of the other authors indicate that one or more separate species may have been confused with the oocysts originally seen by Weidman (1915). Oocysts of these smaller isolates are very similar to reports given for Isospora buriatica Dubey, 1963, the Isospora vulpina Nieschulz and Bos, 1933 isolate studied by Bledsoe (1976), and even Isospora vulpis Galli-Valerio, 1931.

5. Isospora fennechi Prasad, 1961
   1. Host: Fennecus zerda (Fennec fox - type host)
   2. Oocysts: 27.5 x 17.2 (24.5-30.5 x 15-19.5) micrometers.
   3. References: Prasad 1961
   4. Remarks: Prasad (1961) was unable to infect dogs with this coccidian.

6. Isospora neorivolta Dubey and Mahrt, 1978
   1. Synonym: Isospora rivolta of Mahrt, 1967
   2. Synonym: Isospora rivolta of Mahrt, 1968
   3. Host: Canis familiaris (domestic dog - type host)
   4. Oocysts: No measurements provided; stated to be similar to Isospora ohioensis.
   5. References: Dubey and Mahrt 1978; Mahrt 1967, 1968
   6. Remarks: This coccidian is essentially indistinguishable morphologically from Isospora ohioensis. It was described solely on the basis of its developmental biology. Isospora burrowsi develops in he lamina propria of the posterior portion of the small intestine whereas Isospora ohioensis occurs only in the epithelium and throughout the small intestine.

7. Isospora ohioensis Dubey, 1975
   1. Synonym: Cystoisospora ohioensis (Dubey, 1975) Frenkel, 1977
   2. Synonym: Levinea ohioensis (Dubey, 1975) Dubey, 1977
   3. Host: Canis familiaris (domestic dog - type host); experimentally Canis latrans (Coyote); perhaps Vulpes vulpes (Red and silver foxes) and other canids
   4. Oocysts: 24 x 21 (21-27 x 19-23) micrometers. Levine and Ivens (1965) give measurements of what may be Isospora ohioensis (syn. Isospora rivolta (Grassi, 1879) Wenyon, 1923 from the dog) as 23 x 19 (20-27 x 15-24) micrometers. Oocysts that may be Isospora ohioensis (syn. Isospora rivolta of Loveless and Anderson, 1975) from coyotes were measured as 23.9 x 19.2 micrometers by Loveless and Anderson (1975).
   5. References: Dubey 1975a, 1978a, 1978b, 1982; Dubey and Mehlhorn 1978; Dunbar and Foreyt 1985; Heine 1981; Levine and Ivens 1965; Reinhardt 1934; Streitel and Dubey 1976; Wenyon 1923
   6. Remarks: This coccidian is essentially indistinguishable morphologically from both Isospora neorivolta and Isospora burrowsi although it tends to be somewhat larger than Isospora burrowsi. Arthur and Post (1977) reported Isospora ohioensis from coyotes, Canis latrans, but the sizes of the oocysts suggest the species may have been Isospora burrowsi.

8. Isospora pavlodarica Nukerbaeva and Svanbaev, 1973
   1. Host: Alopex lagopus (Arctic or blue fox), Vulpes vulpes (syn. Vulpes fulvus) (Red and silver foxes - type host)
   2. Oocysts: 22.5-25.2 x 19.6-22.4 micrometers.
   3. References: Nukerbaeva and Svanbaev 1973
   4. Remarks: This may be Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi. It may also be a synonym of Isospora vulpis Galli-Valerio, 1931 and/or Isospora vulpina Nieschulz and Bos, 1933.

9. Isospora theileri Yakimoff and Lewkowitsch, 1932
   1. Host: Canis aureus (golden jackal)
   2. Oocysts: 21.2 x 17.1-18.0 micrometers
   3. Reference: Yakimoff and Lewkowitsch, 1932
   4. Remarks: Measurements suggest this may be Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi.

10. Isospora vulpina Nieschulz and Bos, 1933
    1. Synonym: Isospora aprutina Pellerdy, 1974
    2. Synonym: Isospora vulpina var. aprutina (Nieschulz and Bos, 1933) Mantovani, 1965
    3. Synonym: Isospora vulpina var. vulpina (Nieschulz and Bos, 1933) Mantovani, 1965
    4. Host: Vulpes vulpes (syn. Vulpes fulvus) (Red and silver foxes - type host)
    5. Oocysts: 24.9 x 21.4 (21-32 x 19-27) micrometers. Dunlap (1956) gives measurements of 26.7 x 23.1 (21.4-34.4 x 19.0-28.2), Mantovani (1965) 24.5 x 20.6 (22-28 x 18-22), Nukerbaeva and Svanbaev (1973), 25.2 x 19.6 (22.4-28 x 16.8-22.4), Golemansky and Ridzhakov (1975) 25.0 x 20.0 (20.5-30.5 x 16.0-24.0), Bledsoe (1976) 29.7 x 24.3 (25-38 x 21-32), and Glebezdin (1978) 26.2 x 23.6 (22.4-30.8 x 19.6-25.2) micrometers.
    6. References: Bledsoe 1976a, 1976b; Dunlap 1956; Glebezdin 1978; Golemansky and Ridzhakov 1975; Mantovani 1965; Nieschulz and Bos 1933; Nukerbaeva and Svanbaev 1973; Pellerdy 1974; Ridala 1936
    7. Remarks: Bledsoe (1976a, 1976b) was able to transmit oocysts to domestic dogs and it is clear from the photomicrographs that the coccidian he studied was not Isospora canis. Based on the oocyst measurements given by various authors, however, it seems probable that more than one species of coccidia are being confused. It seems likely that some of the coccidia, including the original description, may be referring to Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi. The Isospora vulpina reported by Bledsoe (1976a, 1976b) may represent a synonym of Isospora vulpis Galli-Valerio, 1931, but the original description of the latter species is so poor that is is difficult to tell.

11. Isospora vulpis Galli-Valerio, 1931
    1. Synonym: Eimeria vulpes Patnaik and Acharjy, 1971 lapsus
    2. Hosts: Vulpes bengalensis (Indian fox), Vulpes vulpes (Silver or red foxes)
    3. Oocysts: 25 x 24 micrometers. Svanbaev and Rachmaatullina (1971) measured oocysts as 28.0 x 25.2 (25.2-27.7 x 22.4) and Golemansky and Ridzhakov (1975) 20.0-23.0 x 17.8-21.0 micrometers.
    4. References: Galli-Valerio 1931; Golemansky and Ridzhakov 1975; Patnaik and Acharjy 1971; Svanbaev and Rachmaatullina 1971
    5. Remarks: The measurements provided by Galli-Valerio (1931) suggest that many of the oocysts measured were end-on views resulting in the mean of the measurements coming out to be erroneously more spherical. 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 Bledsae (1976a, 1976b) was actually working with Isospora vulpis rather than Isospora vulpina. However, Galli-Valerio (1931) stated that the oocysts had a micropyle and his measurements suggest subspherical rather than ellipsoidal oocysts. It is likely that he was in error on both accounts and that the description by Svanbaev and Rachmaatullina (1971) is the most accurate.

II. Synonyms, misidentifications, oddities, and species of doubtful validity

1. Cyclospora sp. of Yai, Bauab, Hirschfeld, de Oliveira, and Damaceno, 1997
   1. Host: Canis familiaris (Domestic dog)
   2. Oocysts: No measurements provided
   3. References: Yai et al. 1997
   4. Remarks: The photomicrographs suggest that the oocysts were either Hammondia heydorni or Neospora caninum.

2. Eimeria adleri Yakimoff and Gousseff, 1936
   1. Host: Vulpes velox (Red and silver fox)
   2. Oocysts: Spherical, 21.5 (18.4-24.5) micrometers. Svanbaev (1960) gave measurements of 25.4 x 24.9 (23.6-28.0 x 23.4-27.6) micrometers.
   3. References: Svanbaev 1960; Yakimoff and Gousseff 1934
   4. Remarks: These are likely rodent eimerian pseudoparasites as no valid eimerians are known from Canis spp.

3. Eimeria aurei Bhatia, Chauha, Agrawal, and Ahluwalia, 1979
   1. Host: Canis aureus naria (golden jackal)
   2. Oocysts: 16.3 x 14.7 (15-17 x 13-15) micrometers
   3. Reference: Bhatia et al. 1979
   4. Remarks: This is likely a pseudoparasite as no valid eimerians are known from Canis spp.

4. Eimeria bakanensis Svanbaev and Rachmaatullina, 1971
   1. Host: Vulpes vulpes (Red and silver foxes)
   2. Oocysts: 14.0 x 11.2 (11.0-16.8 x 8.4-11.2) micrometers.
   3. Reference: Svanbaev and Rachmaatullina 1971
   4. Remarks: This is likely a rodent or avian eimerian.

5. Eimeria canis Wenyon, 1923
   1. Host: Canis familiaris (domestic dog)
   2. Oocysts: 18-45 x 11-28 micrometers; Yakimoff and Matschoulsky (1937) give measurements of 24.4 x 12.5 (sic) (21.1-27.4 x 12.6-21.1) and Skofitsch et al. (1983) 37.5 x 23.9 micrometers.
   3. References: Brown and Stammers 1922; Matschoulsky and Tiimofeev 1940; Patnaik 1966; Skofitsch eet al. 1983; Wenyon 1923, 1926; Yakimoff and Matschoulsky 1937
   4. Remarks: These are likely pseudoparasites of rodents, rabbits, and/or birds. Wenyon (1926) thought they resembled rabbit coccidia and the photomicrographs published by Skofitsch et al. (1983) suggest that their isolate was also a rabbit coccidian. No valid eimerians are known from dogs. Skofitsch et al. (1983) were unable to infect dogs with their oocysts.

6. Eimeria heissini Svanbaev, 1956
   1. Synonym: Eimeria hissani Patnaik and Acharjyo, 1971 lapsus
   2. Hosts: Vulpes bengalensis (Indian fox), Vulpes corsac (Corsac fox)
   3. Oocysts: Spherical, 20.4 in diameter.
   4. References: Glebezdin 1978; Patnaik and Acharjyo 1971; Svanbaev 1956
   5. Remarks: These appear to be rodent eimerian pseudoparasites.

7. Eimeria imantauica Nukerbaeva and Svanbaev, 1973
   1. Host: Alopex lagopus (Arctic or blue fox)
   2. Oocysts: 14.0 x 9.8 (12.6-15.4 x 8.4-11.2) micrometers.
   3. References: Nukerbaeva and Svanbaev 1973
   4. Remarks: This appears to be rodent eimerian pseudoparasite.

8. Eimeria li Golemansky, 1975
   1. Host: Vulpes vulpes (Silver or red foxes)
   2. Oocysts: 31.0 x 23.0 (29.0-33.0 x 22.0-25) micrometers
   3. References: Golemansky, 1975; Golemansky and Ridzhakov 1975
   4. Remarks: This appears to be an eimerian pseudoparasite derived either from a rodent or galliform bird

9. Eimeria lomarii Dubey, 1963
   1. Host: Vulpes bengalensis (Indian fox)
   2. Oocysts: 24-29 x 19-22 micrometers
   3. Reference: Dubey 1963
   4. Remarks: This may be a rodent eimerian pseudoparasite. No valid eimerians are known from Vulpes spp.

10. Eimeria lupi Rodionov, 1941
    1. Host: "Lupus vulgaris"
    2. Oocysts: 16.1 x 11.5 micrometers.
    3. Reference: Rodionov 1941
    4. Remarks: No mammal exists with the name "Lupus vulgaris". It is likely that this represents either Canis aureus (Golden jackal) or Vulpes vulpes (Red or silver foxes) as both species have been assigned the trivial name "vulgaris" in the past. It is likely that the coccidian is a pseudoparasite.

11. Eimeria macrotis Mayberry, Bristol, Duszynski, and Reid, 1980
    1. Host: Vulpes velox (syn. Vulpes macrotis neomexicanus) (Kit or swift fox)
    2. Oocysts: 31.6 x 19.1 (28-37 x 17-21) micrometers
    3. Reference: Mayberry et al. 1980
    4. Remarks: This is likely an eimerian pseudoparasite derived from ingesting a reptile such as a lizard.

12. Eimeria mesnili Rastegaieff, 1929
    1. Host: Alopex lagopus (syn. Canis lagopus) (Arctic or blue fox)
    2. Oocysts: 18 x 10.8-14.2 micrometers. In a later paper, Rasteigaieff (1930) reports the measurements as 18 x 10.8-16.2 micrometers.
    3. References: Levine and Ivens 1981; Rastegaieff 1929a, 1930; Ridala 1936
    4. Remarks: This is likely a rabbit coccidian.

13. Eimeria rayi Rao and Bhatavdekar, 1957
    1. Host: Canis familiaris (domestic dog)
    2. Oocysts: 26.8 x 19.8 (21.6-29.1 x 18-21.6) micrometers
    3. Reference: Rao and Bhatavdekar, 1957
    4. Remarks: This is certainly a pseudoparasite. Levine and Ivens (1981) remarked that these coccidia resembled a ruminant coccidian. No valid eimerians are known from dogs.

14. Eimeria sp. of Arthur and Post, 1977
    1. Host: Canis latrans (Coyote)
    2. Oocysts: 9.8 x 9.6 (8.5-11.4 x 8.2-11.2) micrometers
    3. Reference: Arthur and Post 1977
    4. Remarks: Although most certainly a pseudoparasite, the coccidian is very similar to Eimeria zajacae Lindsay, Upton, and Hildreth, 1999 from White-tailed deer, Odocoileus virginianus. The coccidian may have been picked up after it was ingested when the coyote either scavenged a carcass or by coprophagy.

15. Eimeria sp. of Golemansky and Ridzhakov, 1975
    1. Host: Vulpes vulpes (Silver or red foxes)
    2. Oocysts: 33.0 x 20.0 (28.0-38.1 x 17.0-22.5) micrometers.
    3. Reference: Golemansky and Ridzhakov 1975
    4. Remarks: This appears to be an eimerian pseudoparasite derived either from a rodent or a rabbit, as stated by the authors.

16. Eimeria sp. of Poglayen, Guberti, and Leoni, 1985
    1. Host: Vulpes vulpes (Red and silver foxes)
    2. Oocysts: 35.1 x 21.4 micrometers.
    3. Reference: Poglayen et al. 1985
    4. Remarks: This is either a avian, lagomorph, or rodent eimerian

17. Eimeria sp. of Skirnisson, Eydal, Gunnarsson, and Hersteinsson, 1993
    1. Host: Alopex lagopus (Arctic or blue fox)
    2. Oocysts: No measurements provided
    3. Reference: Skirnisson et al. 1993
    4. Remarks: The oocysts were in poor shape and could not be identified. It is likely that they represented a pseudoparasite.

18. Eimeria sp. of Streitel and Dubey, 1976
    1. Host: Canis familiaris (Domestic dog)
    2. Oocysts: No measurements provided
    3. Reference: Streitel and Dubey 1976
    4. Remarks: The authors attempted to inoculate oocysts into a coccidia-free puppy but were unsuccessful. They concluded that the coccidian was most likely a pseudoparasite.

19. Eimeria vulpis Galli-Valerio, 1929
    1. Synonym: Eimeria vulpes Patnaik and Acharjy, 1971, lapsus
    2. Hosts: Vulpes bengalensis (Indian fox), Vulpes vulpes (Red and silver fox)
    3. Oocysts: Svanbaev (1960) gives measurements of 19.5 x 14.5 (18.2-22.0 x 13.0-16.3) and Golemansky and Ridzhakov (1975) 21.9 x 15.9 (17.8-26.0 x 13.0-18.8) micrometers.
    4. References: Galli-Valerio 1929; Golemansky and Ridzhakov 1975; Gousseff 1933; Patnaik and Acharjy 1971; Ridala 1936; Svanbaev 1960; Svanbaev and Rachmatullina 1971
    5. Remarks: These are either a avian, lagomorph, or rodent eimerians

20. Isospora arctopitheci Rodhain, 1933
    1. Synonym: Isospora endocallimici Duszynski and File, 1974
    2. Synonym: Isospora natalensis Elsdon-Dew, 1953
    3. Synonym: Isospora scorzai Arcay-de-Peraza, 1967
    4. Hosts: Natural hosts are Callimico goeldii (Goeld's marmoset), Callithrix penicillata (syn. Hapale jacchus penicillatus) (Black tufted-eared marmoset - type host), Cebus capucinus (Capuchin), perhaps Homo sapiens (Humans), and Saguinus geoffroyi (Red-crested tamarin). Experimental primate hosts include Aotus trivigatus (Night monkey), Ateles fuscipes (Spider monkey), Alouatta pigra (syn. Alouatta villosa) (Howler monkey), and Saimiri sciureus (Squirrel monkey). Experimental non-primate hosts include Canis familiaris (Domestic dog), Nasua nasua (Coatimundi), Potos flavus (Kinkajou), Eiria barbar (Tayra), Felis catus (Domestic cat), and Didelphis marsupialis (Opossum).
    5. Oocysts: 25.5-30.5 x 23.2-25.5 micrometers. A redescription by Hendricks (1974) puts oocysts at 26.0 x 22.9 (20.5-29.5 x 20.5-24.5) micrometers.
    6. References: Duszynski et al. 1999; Hendricks 1974, 1977; Rodhain 1933
    7. Remarks: Originally described from a marmoset, experimental transmission studies have shown this coccidian to have an unsually wide host range covering primates, carnivores, and even marsupials. All reports outside of primates are experimental and it is not known if this parasite actually occurs in non-primates naturally.

21. Isospora bahiensis (de Moura Costa, 1956) Levine, 1978
    1. Synonym: Isospora bigemina var. bahiensis (Stiles, 1891) de Moura Costa, 1956
    2. Host: Canis familiaris (Domestic dog)
    3. Oocysts: 13 x 11.5 (12.3-13.6 x 10.7-12.3) micrometers
    4. Reference: de Moura Costa 1956
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum and the parasite is not a true isosporan.

22. Isospora dutoiti Yakimoff, Matikaschwili, and Rastegaieff, 1933
    1. Eimeria dutoiti Yakimoff, Matikaschwili, and Rastegaieff, 1933, lapsus
    2. Host: Canis aureus (Golden jackal)
    3. Oocysts: 10-13 x 9-11 micrometers
    4. References: Levine and Ivens 1981;Yakimoff et al. 1933
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum and the parasite is not a true isosporan.

23. Isospora sp. (Ashford, 1977) Levine and Ivens, 1981
    1. Synonym: Hammondia sp. of Ashford, 1977
    2. Host: Vulpes vulpes (Silver or red foxes)
    3. Oocysts: 14 x 12 micometers
    4. References: Ashford 1977; Levine and Ivens 1981
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum.

24. Isospora sp. of Dubey, Weisbrode, and Rogers, 1978
    1. Host: Canis familiaris (domestic dog)
    2. Oocysts: No measurements provided of the viable oocysts; only developmental stages within the intestinal tract were studied.
    3. Reference: Dubey et al. 1978
    4. Remarks: This coccidian was found in the lamina propria and, thus, the authors felt that it was probably not Isospora ohioensis. Differences in merozoites structure and site of infection led the authors to conclude that the parasite was not strictly Isospora burrowsi or Isospora neorivolta either. The authors stated that a mixed infection may have been observed.

25. Isospora sp. of Gill, SIngh, Vadehra, and Sethi, 1978
    1. Synonym: Sarcocystis sp. of Gill, SIngh, Vadehra, and Sethi, 1978
    2. Host: Canis familiaris (domestic dog)
    3. Oocysts: 23.3 x 18.2 (17.5-23.8 x 16.3-18.8) micrometers (?).
    4. Reference: Gill et al. 1978
    5. Remarks: The authors fed dogs diaphragm of a water buffalo, Bubalus bubalis, containing cysts of Sarcocystis sp. and received back oocysts which they termed a Sarcocystis sp. Whether or not the dogs became spuriously infected, or whether dormant cytozoites were present in muscle tissue of the buffalo, is unknown. However, the photomicrographs clearly reveal oocysts that are not Sarcocystis and appear most similar to Isospora ohioeneis.

26. Isospora sp. of Golemansky and Ridzhakov, 1975
    1. Synonym: Isospora bigemina (Stiles, 1891) Luhe, 1906 of Golemansky and Ridzhakov, 1975
    2. Host: Vulpes vulpes (Silver or red foxes)
    3. Oocysts: 12.7-17.8 x 10.2-15.2 micrometers.
    4. Reference: Golemansky and Ridzhakov 1975
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum.

27. Isospora sp. of Levine and Ivens, 1965
    1. Synonym: Isospora bigemina (Stiles, 1891) Luhe, 1906 of Levine and Ivens, 1965
    2. Host: Canis familiaris (domestic dog)
    3. Oocysts: 11.7 x 10.4 (10-14 x 10-12) micrometers
    4. Reference: Levine and Ivens 1965
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum.

28. Isospora sp. of Rusvay, 1926
    1. Synonym: Isospora rivolta (Grassi, 1879) Wenyon, 1923 of Rusvay, 1926
    2. Host: Canis familiaris (domestic dog)
    3. Oocysts: 19-23.8 x 16.6-19 micrometers.
    4. Reference: Rusvay 1926
    5. Remarks: This is one of the Isospora ohioensis-like isosporans.

29. Isospora sp. of Rusvay, 1926
    1. Synonym: Isospora bigemina of Rusvay, 1926 (small form oocysts)
    2. Host: Canis familiaris (domestic dog)
    3. Oocysts: 11.9-14.3 x 8.9-11.9 micrometers.
    4. Reference: Rusvay 1926
    5. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum.

30. Isospora sp. of Skirnisson, Eydal, Gunnarsson, and Hersteinsson, 1993
    1. Host: Alopex lagopus (Arctic or blue fox)
    2. Oocysts: No measurements provided
    3. Reference: Skirnisson et al. 1993
    4. Remarks: The oocysts were in poor shape and could not be identified.

31. Isospora sp. of Svanbaev, 1956
    1. Synonym: Isospora buriatica Dubey, 1963 of Svanbaev, 1956 pro parte
    2. Host: Vulpes corsac (Corsac fox)
    3. Oocysts: 25.8 x 19.0 micrometers.
    4. Reference: Svanbaev 1956
    5. Remarks: Although perhaps a separate species, this may in fact represent Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi.

32. Isospora sp. of Thornton, Bell, and Reardon, 1974
    1. Synonym: Isospora rivolta of Thornton, Bell, and Reardon, 1974
    2. Host: Canis latrans (coyote)
    3. Oocysts: No measurements provided
    4. References: Thornton et al. 1974
    5. Remarks: This is likely Isospora ohioensis, Isospora neorivolta, or Isospora burrowsi.

33. Isospora triffitti Nukerbaeva and Svanbaev, 1973
    1. Host: Alopex lagopus (Arctic or blue fox), Vulpes vulpes (syn. Vulpes fulvus) (Red and silver foxes)
    2. Oocysts: 11.2-12.6 x 9.8-11.2 micrometers
    3. References: Nukerbaeva and Svanbaev 1973
    4. Remarks: It is likely that the oocysts were either Hammondia heydorni or Neospora caninum and the parasite is not a true isosporan.

34. Isospora tropicalis Mukherjea and Krassner, 1965
    1. Host: Canis aureus (Golden jackal)
    2. Oocysts: Average 16 micrometers in diameter
    3. Reference: Mukherjea and Krassner 1965
    4. Remarks: The photomicrographs clearly reveal this to be a Sarcocystis sp.

35. Isospora uralense Rodionov, 1941
    1. Host: "Lupus vulgaris"
    2. Oocysts: Spherical, 11.5 micrometers in diameter.
    3. Reference: Rodionov 1941
    4. Remarks: No mammal exists with the name "Lupus vulgaris". It is likely that this represents either Canis aureus (Golden jackal) or Vulpes vulpes (Red or silver foxes) as both species have been assigned the trivial name "vulgaris" in the past. The coccidian represents either Hammondia heydorni or Neospora caninum and is not a true isosporan.

36. Isospora wetzeli Sugar, 1980
    1. Host: Vulpes vulpes (Red and silver foxes)
    2. Oocysts: 13-16.6 x 15-19 micrometers
    3. Reference: Sugar 1980
    4. Remarks: This is a Sarcocystis sp.

III. Tissue cyst forming coccidia often confused with canid Isospora spp.

1. Hammondia heydorni (Tadros and Laarman, 1983) Dubey, 1977
   1. non Isospora wallacei Tadros and Laarman, 1983
   2. Synonym: Isospora bigemina (Stiles, 1891) Luhe, 1906 "small form" of various authors, pro parte
   3. Synonym: Isospora bigemina of Dubey and Fayer, 1976
   4. Synonym: Isospora wallacei Dubey, 1976
   5. Definitive hosts: Canis familiaris (domestic dog - type host); experimentally Canis latrans (Coyote); perhaps Mustela vison (Mink)
   6. Oocysts: Dubey and Fayer (1976) reported oocysts as spherical, 12 x 11 micrometers. Nassar et al. (1983) gives 11.9 x 11.1 (10.0-14.5 x 9.3-13.1) and Shankar et al. (1991) 11.0-14.0 x 10.5-13.0 micrometers.
   7. References: Blagburn et al. 1988; Dubey 1977, 1982; Dubey and Fayer 1976; Dubey and Williams 1980; Heydorn 1973; Istvan and Mikaly 1988; Levine and Ivens 1981; Matsui et al. 1981, 1986, 1987; Nassar et al. 1983; Shankar et al. 1991
   8. Remarks: This coccidian is obligatorily heteroxenous and uses a wide range of intermediate hosts where muscle tissue cysts are formed. Intermediate hosts include ruminants, cervids, and rodents. Over the years, the most commonly employed term has been the "small form" of Isospora bigemina (Stiles, 1891) Luhe, 1906. Oocysts are passed unsporulated.

2. Neospora caninum Dubey, Carpenter, Speer, Topper, and Uggla, 1988
   1. Synonym: Isospora bigemina (Stiles, 1891) Luhe, 1906 "small form" of various authors, pro parte
   2. Definitive host: Canis familiaris (domestic dog)
   3. Oocysts: Spherical to subspherical, 11.7 x 11.3 (10.6-12.4 x 10.6-12.0) micrometers
   4. References: Anderson et al. 1991; Dubey 1992, 1999a, 1999b; Dubey and Lindsay 1989, 1996; Dubey et al. 1988a, 1988b, 1990; Lindsay and Dubey 1989; Lindsay et al. 1999; McAllister et al. 1998
   5. Remarks: Oocysts are passed unsporulated. Neospora caninum is a causitive agent of canine encephalomyelitis and myositis, resulting in hindlimb paralysis and often death. To date, it has not yet been determined whether this coccidian is obligatorily or facultatively heteroxenous. If oocysts inoculated into dogs results in oocysts being passed in the feces, then this could be considered just another species of Toxoplasma. If oocysts inoculated into dogs fail to result in oocyst production, then this could be regarded as another species of Hammondia. And, if like some people you believe that the genus Hammondia should be synonymized with Toxoplasma, then you still end up with another species of Toxoplasma.

I. Named and potentially valid felid Isospora spp.

1. Isospora felis Wenyon, 1923
   1. Synonym: Cystoisospora felis (Wenyon, 1923) Frenkel, 1977
   2. Synonym: Diplospora bigemina of Wasielewski, 1904 pro parte
   3. Synonym: Isospora bengalensi Patnaik and Acharjyo, 1971
   4. Synonym: Isospora bigemina of Swellengrebel, 1914
   5. Synonym: Isospora cati Marotel, 1921
   6. Synonym: Isospora felis var. servalis Mackinnon and Dibb, 1938
   7. Synonym: Isospora leopardi Agrawal, Ahluwalia, Bhatia, and Chauhan, 1981
   8. Synonym: Isospora nandankani Patnaik and Acharjyo, 1977
   9. Synonym: Isospora pantheri Agrawal, Ahluwalia, Bhatia, and Chauhan, 1981
   10. Synonym: Isospora pardusi Patnaik and Acharjyo, 1971
   11. Synonym: Isospora rivolta of Dobell and Connor, 1921
   12. Synonym: Levinea felis (Wenyon, 1923) Dubey, 1977
   13. Synonym: Lucetina felis (Wenyon, 1923) Henry and Leblois, 1926
   14. Hosts: Felis catus (Domestic cat - type host), Felis silvestris (European wild cat), Leopardus pardalis (syn. Felis pardalis) (Ocelot), Leptailurus serval (syn. Felis serval) (Serval), Lynx lynx (Eurasian lynx), Neofelis nebulosa (Clouded leopard), Panthera leo (syn. Leo leo) (Lion), Panthera onca (syn. Leo onca) (Jaguar), Panthera pardus (Leopard), Panthera tigris (syn. Leo tigris) (Tiger), Prionalilurus bengalensis (syn. Felis bengalensis) (Leopard cat)
   15. Oocysts: Swellengrebel (1914) gave measurements of 39-47 x 26-37, Wenyon (1923) 45 x 33 (39-48 x 26-37), Nieschulz (1925) 42.4 x 33.4 (39-44 x 32.36), Sphren and Cramer (1931) 35-45 x 23-35, Matschoulsky and Tiimofeev (1940) 37.7 x 30.8 (33.6-44.1 x 27.3-35.7), Tomimura (1957) 43.5 x 32.9 (38-49 x 28-37), Costa and Freitas (1959) 32.8-44.4 x 27.0-33.6, Lickfeld (1959) 43 x 32 (32-53 x 27-43), Shah (1970b) 41.6 x 30.5 (38-51 x 27-39), de Rocha and Lopes (1971) 39.4 x 30.5 (32.1-46.5 x 26.2-37.2), Gonzalez Castro et al. (1972) 41.3 x 32.2, and Ng'ang'a et al. (1994) 42.4 x 34.0 micrometers. Chaudhuri and Choudury (1980) reported oocysts from a "tigon" as 46.0 x 34.0 (40.8-49.3 x 28.9-34.0), Chaudhuri and Choudury (1982) from a tiger 46.0 x 31.0 (41.0-49.5 x 29.0-34.2), Yakimoff et al. (1933) from a lion as 44.5 x 33.6 (37.8-50.4 x 28.8-36), Yakimoff et al. (1933) from a tiger 39.5 x 30.5 (34-45 x 22.2-35.5), and Yakimoff et al. (1933) from one lynx as 45.5 x 35 (39.5-50.4 x 30.6-39.6) and from another lynx as 36 x 29 (30-42 x 24.5-33) micrometers. Bjork et al. (2000) measured oocysts from a lion as 43.0 x 30.0 (39.0-45.0 x 25.0-35.0) micrometers.
   16. References: Agrawal et al. 1981; Bjork et al. 2000; Chaudhuri and Chaoudhury 1982; Costa and Freitas 1959; Daly and markus 1981; Dubey 1973; Dubey and Frenkel 1972a; Fayer and Frenkel 1979; Ferguson et al. 1980; Frenkel and Dubey 1972; Gonzalez Castro et al. 1972; Heine 1981; Hitchcock 1955; Lickfeld 1959; MacKenna and Charleston 1982; Matschoulsky and Tiimofeev 1940; Mehlhorn and Markus 1976; Ng'ang'a et al. 1994; Nieschulz 1925; Patnaik and Acharjyo 1971, 1977; de Rocha and Lopes 1971; Shah 1970a, 1970b, 1971; Sphren and Cramer 1931; Swellengrebel 1914; Tomimura 1957; Wenyon 1923; Yakimoff et al. 1933
   17. Remarks: The only difference between Isospora bengalensi Patnaik and Acharjyo, 1971 from Prionalilurus bengalensis (syn. Felis bengalensis) and Isospora felis are the sporocyst measurements. Based on the line drawing, it is apparent that the measurements given by Patnaik and Acharjyo (1971) represent a printing error and that the sporocysts are actually larger and more similar to Isospora felis.

2. Isospora leonina Mandal and Ray, 1960
   1. Host: Panthera leo (syn. Felis leo) (Lion)
   2. Oocysts: 30-32 x 28-31 micrometers. Patnaik and Acharjyo (1970) give measurements of 31.8 x 29 (30-33 x 28-30) micrometers.
   3. References: Mandal and Ray, 1960; Patnaik and Acharjyo 1970
   4. Remarks: This appears to be a carnivore isosporan, but whether or not this is a true species of the lion or a pseudoparasite is not yet known.

3. Isospora rivolta (Grasse, 1879) Wenyon, 1923
   1. Synonym: Cystoisospora frenkeli Arcay, 1981
   2. Synonym: Cystoisospora rivolta (Grassi, 1879) Frenkel, 1977
   3. Synonym: Coccidium rivolta Grassi, 1879
   4. Synonym: Coccidium rivoltae Leuckart, 1996 pro parte
   5. Synonym: Diplospora bigemina Wasielewski, 1904 pro parte
   6. Synonym: Isospora felina Patnaik and Acharjyo, 1971
   7. Synonym: Isospora frenkeli Arcay-de-Peraza, 1976
   8. Synonym: Isospora mohini Agrawal, Ahluwalia, Bhatia, and Chauhan, 1981
   9. Synonym: Isospora novocati Pellerdy, 1974
   10. Synonym: Isospora rivoltae Dobell, 1919
   11. Synonym: Isospora sp. of Pande, Bhatia, Chauhan, and Garg, 1970
   12. Synonym: Levinea rivolta (Grassi, 1879) Dubey, 1977
   13. Synonym: Lucetina rivolta (Grassi, 1879) Henry and Leblois, 1926
   14. Hosts: Acinonyx jubatus (Cheetah), Felis catus (Domestic cat - type host), Felis chaus (Jungle cat), Felis silvestris (European wild cat), Neofelis nebulosa (Clouded leopard), Panthera pardus (Leo pardus) (Leopard), Panthera leo (Lion), Panthera tigris (syn. Leo tigris) (Tiger), Prionalilurus bengalensis (syn. Felis bengalensis) (Leopard cat)
   15. Oocysts: Nieschulz (1925) reported oocysts as 25.5 x 22.5 (22-28 x 18-25), Sphren and Cramer (1931) 20-28 x 18-22, Matschoulsky and Tiimofeev (1940) 26.4 x 21.6 (21.0-29.4 x 18.9-25.2), Costa and Freitas (1959) 22.8 x 19.5 (19.8-23.9 x 18.3-20.6), Dubey and Pande (1963) 28 x 23 (27-30 x 20-25), Shah (1970a) 25.0 x 21.1 (21-28 x 18-23), de Rocha and Lopes (1971) 26.3 x 21.8 (21.9-30.0 x 18.6-26.0), Pellerdy (1974) 27 (24-30) for spherical forms and 26 x 24 (22-30 x 21-27) for subspherical forms, Dubey (1979) 25.4 x 23.4 (23-29 x 20-26), and Ng'ang'a et al. (1994) 25.2 x 23.2 micrometers. Yakimoff et al. (1933) measured an isolate from tigers as 22.1 x 16.5 (18-25 x 13-20), one from a leopard as 23 x 17.5 (20-25.5 x 15-21), and one from a jungle cat as 23.4-27 x 18-23 micrometers. Bjork et al. (2000) measured oocysts from a lion as 25.0 x 21.0 (30.0-20.0 x 25.0-15.0) micrometers.
   16. References: Agrawal et al. 1981; Arcay 1981; Bjork et al. 2000; Brosigke et al. 1982; Costa and Freitas 1959; Dubey 1979; Dubey and Frenkel 1972a; Dubey and Pande 1963; Fayer and Frenkel 1979; Frenkel and Dubey 1972; Heine 1981; MacKenna and Charleston 1982; Matsui et al. 1993a, 1993b; Ng'ang'a et al. 1994; Nieschulz 1925; Patnaik and Acharjyo 1971; Pellerdy 1974; Penzhorn et al. 1994; de Rocha and Lopes 1971; Shah 1970a; Sphren and Cramer 1931; Yakimoff et al. 1933
   17. Remarks: The sporocyst length measurement of Isospora felina Patnaik and Acharjyo 1971 from Prionalilurus bengalensis (syn. Felis bengalensis) (Leopard cat) represents the only distinction between this species and Isospora rivolta. Based on the line drawing, however, it appears that the given measurement represents a printing error and that this coccidian is probably not a distinct species.

II. Synonyms, misidentifications, oddities, and species of doubtful validity

1. Eimeria anekalensis Rajasekariah, Hegde, Srinivasa Gowda, Abdul Rahman, and Subbarao, 1971
   1. Host: Panthera pardus (Felis pardus) (Leopard)
   2. Oocysts: 26.5 x 19.8 (22-30 x 18-22) micrometers
   3. Reference: Rajesekariah et al. 1971
   4. Remarks: This is likely an avian pseudoparasite.

2. Eimeria cati Yakimoff, 1932
   1. Host: Felis catus (Domestic cat - type host), Felis chaus (Jungle cat), Felis silvestris (European wild cat)
   2. Oocysts: Oocysts were originally reported as spherical to ovoidal, 20.8 x 19.9 (18-23.4 x 14.4-19.8) micrometers. Later, Yakimoff (1933) reported spherical, ovoidal, and ellipsoidal forms. Spherical oocysts measured 16.2-21.6 micrometers in diameter, ovoidal oocysts 19.8 x 16.2 (18.9-23.4 x 14.4-19.8), and ellipsoidal oocysts 20.8 x 17.1 (18.0-23.4 x 14.4-19.8) micrometers. Dubey and Pande (1963) reported oocysts in the jungle cat as spherical to subspherical, 21 x 19 (19-25 x 19-22) micrometers.
   3. References: Dubey and Pande 1963; Matschoulsky and Tiimofeev 1940; Mirza 1970; Yakimoff 1932, 1933
   4. Remarks: These appear to be rodent eimerian pseudoparasites.

3. Eimeria chaus Rysavy, 1954
   1. Host: Felis chaus (syn. Chaus chaus) (Jungle cat)
   2. Oocysts: 20.4 x 18 (18.3-24.4 x 14.3-22.4) micrometers
   3. Reference: Rysavy 1954
   4. Remarks: This is either a rodent or avian coccidian.

4. Eimeria felina Nieschulz, 1924
   1. Host: Felis catus (Domestic cat - type host), Felis chaus (Jungle cat), Panthera leo (Lion)
   2. Oocysts: 24 x 14.5 (21-26 x 13-17) micrometers. Rastegaieff (1929) gives measurements as 20.7 x 16.2 and Dubey and Pande (1963) 19 x 15 (15-19 x 11-17) micrometers.
   3. References: Bjork et al. 2000; Dubey and Pande 1924; Fameree and Cotteleer 1976; Nieschulz 1924; Patnaik 1966; Rastegaieff 1929b, 1930
   4. Remarks: These appear to be rodent or avian eimerian pseudoparasites.

5. Eimeria hammondi Dubey and Pande, 1963
   1. Host: Felis chaus (Jungle cat)
   2. Oocysts: 24-29 x 19-22 micrometers.
   3. Reference: Dubey and Pande 1963
   4. Remarks: This appears to be a rodent or avian eimerian pseudoparasite.

6. Eimeria (?) hartmanni Rastegaieff, 1930
   1. Hosts: Panthera pardus (Felis pardus) (Leopard)
   2. Oocysts: Unsporulated oocysts 22.8 x 14.4 micrometers. Rajesekariah et al. (1971) reported oocysts as 21.5 x 16.5 (20-22.5 x 14-19) micrometers.
   3. References: Rastegaieff 1930; Rajesekariah et al. 1971
   4. Remarks: The original description was based on unsporulated oocysts. Levine and Ivens (1981) thought this might be a rabbit coccidian.

7. Eimeria lyncis Anpilogova and Sokov, 1973
   1. Host: Lynx lynx (Eurasian lynx)
   2. Oocysts: 33.3 x 22.8 (29.7-37.8 x 18.9-27.0) micrometers
   3. Reference: Anpilogova and Sokov, 1973
   4. Remarks: It is likely that this is either a rodent or avian eimerian.

8. Eimeria mathurai Dubey and Pande, 1963
   1. Host: Felis chaus (Jungle cat)
   2. Oocysts: 22 x 19 (20-28 x 16-20) micrometers.
   3. Reference: Dubey and Pande 1963
   4. Remarks: This appears to be a rodent or avian eimerian pseudoparasite.

9. Eimeria novowenyoni Rastegaieff, 1929
   1. Host: Panthera pardus (Felis pardus) (Leopard), Panthera tigris (syn. Felis tigris) (Tiger - type host)
   2. Oocysts: Spherical, 14-15 micrometers in diameter. Rajesekariah et al. (1971) reported oocysts as 18-20 micrometers in diameter.
   3. References: Rajesekariah et al. 1971; Rastegaieff 1929, 1930

10. Eimeria sp. of Anpilogova and Sokov, 1973
    1. Host: Lynx lynx (Eurasian lynx)
    2. Oocysts: 40.5 x 27.0 micrometers
    3. Reference: Anpilogova and Sokov 1973
    4. Remarks: It is likely that this is either a rodent or avian eimerian.

11. Eimeria tadzhikistanica Anpilogova and Sokov, 1973
    1. Host: Lynx lynx (Eurasian lynx)
    2. Oocysts: 31.2 x 23.5 (24.3-32.4 x 18.9-27.0) micrometers
    3. Reference: Anpilogova and Sokov 1973
    4. Remarks: It is likely that this is either a rodent or avian eimerian.

12. Isospora sp. of Bjork, Averbeck, and Stromberg, 2000
    1. Host: Panthera leo (Lion)
    2. Oocysts: Spherical, 15.0 in diameter
    3. Reference: Bjork et al. 2000
    4. Remarks: It is likely that this is either a Besnoitia sp., Hammondia hammondi, or Toxoplasma gondii.

III. Tissue cyst forming coccidia often confused with felid Isospora spp.

1. Besnoitia besnoiti (Marotel, 1912) Henry, 1913
   1. Synonym: Gastrocystis besnoiti (Marotel, 1912) Brumpt, 1913
   2. Synonym: Gastrocystis robini Brumpt, 1913
   3. Synonym: Globidium besnoti (Marotel, 1912) Wenyon, 1926
   4. Synonym: Isospora besnoiti (Marotel, 1912) Tadros and Laarman, 1976
   5. Synonym: Sarcocystis besnoiti Marotel, 1913
   6. Definitive host: Tentatively, Felis catus (Domestic cat - type species), Felis silvestris (syn. Felis libyca) (Common wild cat)
   7. Intermediate hosts: A wide variety of ruminants and rodents can be infected.
   8. Oocysts: Tentatively, 14.2-16.0 x 11.6-14.2 micrometers
   9. References: Diesing et al. 1988; Peteshev et al. 1974; Rommel, 1975
   10. Remarks: Peteshev et al. (1974) originally reported cats to shed oocysts after eating cysts isolated from cattle. However, neither Rommel (1975) nor Diesing et al. (1988) were able to repeat these studies.

2. Besnoitia darlingi (Brumpt, 1913) Mandour, 1965
   1. Synonym: Besnoitia panamensis Schneider, 1965
   2. Synonym: Besnoitia saurina Garnham, 1966
   3. Synonym: Fibrocystis darlingi (Brumpt, 1913) Babudieri, 1932
   4. Synonym: Sarcocystis darlingi Brumpt, 1913
   5. Definitive host: Felis catus (Domestic cat)
   6. Intermediate hosts: Lizards of the genus Ameiva (Teiidae) and Basiliscus (Corytophanidae), Didelphis marsupialis (Opossum) and a variety of other marsupials; experimentally a variety of rodents, and perhaps Carollia perspicillaata (Short-tailed bat).
   7. Oocysts: Smith and Frenkel (1977) reported sporulated oocysts of the opossum/cat cycle as 12.0 x 10.3 (11.2-12.8 x 9.6-11.2) and unsporulated oocysts as 12.3 x 11.9 (11.2-12.8 x 10.8-12.8) micrometers.
   8. References: Schneider 1965, 1966, 1967a, 1967b, 1967c; Smith and Frenkel 1977; Stabler and Welch 1961
   9. Remarks: Cross-imunity studies by Schneider (1967c) suggested that the species occurring in opossums and lizards may be synonyms; however, this is still unclear.

3. Besnoitia wallacei (Tadros and Laarman, 1976) Frenkel, 1977
   1. non Isospora wallacei Dubey, 1976
   2. Synonym: Besnoitia sp. Wallace and Frenkel, 1975
   3. Synonym: Isospora bigemina "large type" of Ito, Tsunoda, Nishikawa, and Matsui, 1974
   4. Synonym: Isospora bigemina "large type" of Ito, Tsunoda, Nishikawa, and Matsui, 1978
   5. Synonym: Isospora wallacei Tadros and Laarman, 1976
   6. Definitive host: Felis catus (Domestic cat - type host)
   7. Intermediate hosts: Tissue cysts occur in a variety of rodents and, experimentally, in rabbits. Guinea pigs are not susceptible.
   8. Oocysts: Unsporulated oocysts spherical, 16 x 13 (15-18 x 12-15) micrometers. Sporulated oocysts 17 x 12 (16-19 x 10-13) micrometers. Ito et al. (1974, 1978) gave measurements of 15.9 x 13.1 (14.0-17.8 x 12.0-15.0), McKenna and Charleston (1980), 16.9 x 14.6 (14.7-18.0 x 13.0-16.3), and Ng'ang'a et al. (1994) 16.6 x 14.5 (14.0-19.4 x 12.6-17.8) micrometers.
   9. References: Frenkel 1977; Ito and Shimura 1986; Ito et al. 1974, 1978; McKenna and Charleston 1980; Ng'ang'a et al. 1994; Wallace and Frenkel 1975

4. Hammondia hammondi Frenkel, 1974
   1. Synonym: Isospora bigemina (Stiles, 1891) Luhe, 1906 "small form" of various authors, pro parte
   2. Synonym: Isospora datusi Overdulve, 1978
   3. Synonym: Isospora hammondi (Frenkel, 1974) Tadros and Laarman, 1976
   4. Synonym: Toxoplasma hammondi (Frenkel, 1974) Levine, 1977
   5. Definitive host: Felis catus (Domestic cat - type host), Felis silvestris (European wild cat)
   6. Intermediate hosts: Rodents, rabbits, primates, canids, ruminants and swine have all been shown to be capable of carrying infective asexual stages
   7. Oocysts: Unsporulated oocysts 11.4 x 10.6 (1.2-13.2 x 10.5-12.5) micrometers. Sporulated oocysts 13.2 x 10.6 (12.6-13.8 x 10.0-10.7) micrometers. Shimura and Ito (1986) gave measurements of 12.7 x 11.5 (11.0-14.0 x 9.5-13.0) for unsporulated and 13.0 x 11.4 (11.5-14.0 x 9.5-13.0) micrometers for sporulated oocysts, and Rommel and Seyerl (1976) reported sporulated oocysts as 13.8 x 12.2 (11.7-17.5 x 10.2-14.6) micrometers.
   8. References: Dubey 1975b; Dubey and Streitel 1976; Dubey et al. 1978; Eydelloth 1977; Frenkel 1974; Frenkel and Dubey 1975; Markus 1979; Mason 1978; Rommel and Seyerl 1976; Shimura and Ito 1986, 1987; Wallace 1975
   9. Remarks: This coccidian is obligatorily heteroxenous, using felids as definitive hosts and rodents as intermediate hosts. Cysts develop primarily within skeletal muscle, although other sites are infected as well.

5. Hammondia pardalis Hendricks, Ernst, Courtney, and Speer, 1979
   1. Synonym: Isospora sp. of Long and Speer, 1977
   2. Synonym: Toxoplasma pardalis (Hendricks, Ernst, Courtney, and Speer, 1979) Levine and Ivens, 1981
   3. Definitive hosts: Felis catus (Domestic cat), Felis pardalis (Ocelot - type host), Herpailurus yaguarondi (syn. Felis yaguarondi) (Jaguarondi); perhaps Panthera onca (Jaguar) and Puma concolor (syn. Felix concolor) (Puma)
   4. Intermediate hosts: Rodents
   5. Oocysts: 40.8 x 28.5 (36-46 x 25-35) micrometers.
   6. References: Abbitt et al. 1993; Hendricks et al. 1979; Patton et al. 1986
   7. Remarks: This coccidian is obligatorily heteroxenous, using felids as definitive hosts and rodents as intermediate hosts where cysts within skeletal muscle develop. The differential diagnosis for this species is Isospora felis Wenyon, 1923, which has no micropyle whereas oocysts of Hammondia pardalis possess suture/micropyle-like structures at each end. Studies by Abbitt et al. (1993) have suggested that this parasite may cause protozoal abortions in cattle.

6. Toxoplasma gondii (Nicholle and Manceaux, 1908) Nicholle and Manceaux, 1909
   1. Synonym: Haemogregarina francae de Mello, 1915
   2. Synonym: Leishmania gondii Nicholle and Manceaux, 1908
   3. Synonym: Toxoplasma canis de Mello, 1910
   4. Synonym: Toxoplasma caviae Carini and Migliano, 1916
   5. Synonym: Toxoplasma columbae Yakimoff and Kohl-Yakimoff, 1912
   6. Synonym: Toxoplasma crociduri Galli-Valerio, 1933
   7. Synonym: Toxoplasma cuniculi (Splendore, 1908)
   8. Synonym: Toxoplasma francae (de Mello, 1915) Wenyon, 1926
   9. Synonym: Toxoplasma fulicae de Mello, 1935
   10. Synonym: Toxoplasma gallinarum Hepding, 1939
   11. Synonym: Toxoplasma hominis Wolf, Cowen, and Paige, 1939
   12. Synonym: Toxoplasma laidlawi Coutelen, 1932
   13. Synonym: Toxoplasma musculi Sangiorgi, 1913
   14. Synonym: Toxoplasma nikanorovi Zasukhin and Gaisky, 1930
   15. Synonym: Toxoplasma pyrogenes Castellani, 1913
   16. Synonym: Toxoplasma ratti Sangiorgi, 1915
   17. Synonym: Toxoplasma sciuri Coles, 1914
   18. Synonym: Toxoplasma talpae von Prowazek, 1910
   19. Synonym: Toxoplasma wenyoni Coutelen, 1932
   20. Definitive hosts: Felis catus (Domestic cat) and numerous other felids
   21. Intermediate hosts: Over 350 different species of mammals and birds have been shown to be suitable intermediate hosts
   22. Oocysts: Overdulve (1970c) reported unsporulated oocysts as 11.2 x 10.4 (10.3-12.1 x 9.7-10.9) and Dubey et al. (1970) 12 x 10 (11-13 x 9-11) micromters. Dubey et al. (1970b) gave measurements of sporulated oocysts as 12.5 x 11 (11-14 x 10-11), Ng'ang'a et al. (1994) 12.0 x 10.3, and Lindsay et al. (1997) 13 x 11 micrometers.
   23. References: Davis and Dubey 1995; Dubey 1996; Dubey and Frenkel 1972b, 1973, 1976, 1998; Dubey et al. 1970a, 1970b, 1991, 1995, 1996, 1997a, 1997b, 1998; Eyles and Frenkel 1952; Frenkel et al. 1969, 1970; Lindsay et al. 1997; Miller et al. 1972; Ng'ang'a et al. 1994; Overdulve 1970a, 1970b, 1970c, 1978; Pfefferkorn et al. 1977; Sheffield and Melton 1968, 1970; Siim et al. 1969

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