New Genera and Higher Taxa of Septate Gregarines (Protozoa,Apicomplexa)*

SYNOPSIS The following new taxa of septate gregarines are established: METAMERIDAE fam. nov. includes genera Metamera Duke, Deutomera Bhatia & Setna, and Cognettia gen. n., the last with the type and only species C. legeri (Cognetti de Martiis); family GREGARINIDAE Labbé includes Degiustia gen. n. with the type and only species D. hyallela (Batten & DeGiusti); Cirrigregarina gen. n., with the type species C. spissa (Henry) and another species, C. kamenote (Hoshide); and Molluskocystis gen. n., with the type and only species M. pterotracheae (Stuart). Family HIRMOCYSTIDAE (Grassé) includes Arachnocystis gen. n., with the type species A. arachnoidea (Devdhar & Gourishankar) and 3 other species A. nitida (Geus), A. oribatidae (Geus), A. scutovertexi (Erhardová). Family ACTINOCEPHALIDAE (Léger), subfamily Actinocephalinae (Léger), includes Chilogregarina gen. n., with the type species C. striata (Léger & Duboscq) and 3 other species C. brasiliensis (Pinto), C. dujardini (Schneider), and C. stella (Léger).     

Taxonomy and Review of the Coccidian Genus Cryptosporidium (Protozoa,Apicomplexa)1

Reports of Cryptosporidium in various hosts and cross-transmission experiments are reviewed. Cryptosporidium has been found in mammals (Primates, Artiodactyla, Perissodactyla, Carnivora, Lagomorpha, and Rodentia), birds, reptiles, and fish. The only cross-transmission attempts that have been made have been from mammals to other mammals and to a few birds. Names have been given to 19 “species,” but it is concluded that only four of these should be considered valid at present. These are: C. muris Tyzzer, 1907 in mammals, C. meleagridis Slavin, 1955 in birds, C. crotali Triffit, 1925 in reptiles, and C. nasorum Hoover, Hoerr, Carlton, Hinsman & Ferguson, 1981 in fish.     

The taxonomy of Sarcocystis (Protozoa, Apicomplexa) species

The taxonomy of the heteroxenous apicomplexan protozoan genus Sarcocystis was reviewed, and a list of 122 species with their synonyms and hosts given. Both definitive and intermediate hosts are known for only 56 species. The fine structure of the sarcocyst wall may change with age and is not considered necessarily satisfactory for separating species. Specificity for the intermediate host is not narrow for all species. Earlier work on transmission of the parasite from one intermediate host to another should be repeated in the light of present knowledge of the life cycle of species of Sarcocystis.     

A Check-List of the Species of the Genus Leucocytozoon (Apicomplexa,Plasmodiidae)*

SYNOPSIS. A list is given of 67 named species of Leucocytozoon considered valid at present. Another list is given of 1 reptilian and 146 avian hosts of named species of Leucocytozoon. A 3rd list is given of 34 synonyms. Leucocytozoon has been named from about 0.7% of the approximately 8600 species of birds that exist. The following new taxonomic names are included: L. danilewskyi var. hirundinis Sergent & Sergent, 1905 is emended to L. hirundinis; Saurocytozoon tupinambi Lainson & Shaw, 1969 is changed to Leucocytozoon (Saurocytozoon) tupinambi (Lainson & Shaw, 1969).     

A Check-List of the Species of the Genus Haemoproteus (Apicomplexa,Plasmodiidae)*

SYNOPSIS. A list of the presumably valid named species of the genus Haemoproteus Kruse, 1890 (Apicomplexa, Plasmodiidae) is given. It includes 133 species, 5 varieties and 1 subspecies. One new species is named: H. serini n. sp. for the form in the canary Serinus canarius called H. zuenyoni by Sergent & Sergent (90). The following emendations are made: H. aegyptius for H. cellii var. aegyptius Mohammed, 1958; H. cellii for H. noctuae var. cellii Coatney & Roudabush, 1937; H. hirundinis for H. danilewskii var. hirundinis Sergent & Sergent, 1905; H. tinnunculus for H. danilewskyi var. tinnunculus Wasielewski & Wiilker, 1918; H. crumenius for H. crumenium (Hirst, 1905) Coatney, 1936; H. mathislegeri for H. herodiadis var. mathislegeri Mohammed, 1958; H. nebraskensis for H. noctuae var. nebraskensis Coatney & Roudabush, 1937; H. rileyi for Hoemoproteus rileyi Malkani, 1936; H. sauianae for H. sd-vianae Tendeiro, 1947; H. asturisdussumieri for H. asturis-dussumieri de Mello, 1935 emend. de Mello, 1937; H. xantholemae for H. xantholaemi Zargar, 1945. Of the 133 named species, 114 occur in birds, 16 in reptiles and 3 in amphibia; 14 orders and 47 families of birds are represented. The generic names Halteridium LabbC, 1894, Haemocystidium Castellani & Willey, 1904 and Simondia Garnham, 1966 are considered synonyms of Haemoproteus.     

Cross-transmission of Eimeria spp. (Protozoa, Apicomplexa) of rodents--a review

A total of 169 cross-transmission attempts has been made with 44 (11.8%) of the 372 named species of Eimeria of rodents. Of these, 161 were rodent-to-rodent, 6 rodent-to-lagomorph, and 1 each rodent-to-carnivore and rodent-to-bird. None of the last three categories was successful. In the rodent-to-rodent combinations, 39 (80%) of the 49 attempts to transmit a coccidian species from one rodent species to another of the same genus were successful, and only 14 (12.5%) of the 112 attempts to transmit a coccidium to a rodent of a different genus were successful. Eight of the successful attempts were with E. chinchillae, which was the only truly euryxenous species of Eimeria in the group. Two successful attempts were between between the closely related rodent genera Spermophilus and Cynomys, and two were both of E. separata from Rattus norvegicus to some genetic strains but not to others of Mus musculus. One attempt with E. vermiformis from Mus musculus to Rattus norvegicus required treatment of the rat with the immunosuppressant dexamethasone to succeed. More cross-transmission studies are needed to determine the host spectra of the species of Eimeria and other coccidian genera, and to determine the roles of genetics and immunosuppression in their transmission.     

Progress in Taxonomy of the Apicomplexan Protozoa

In 1987, 4516 species and 339 genera of the phylum Apicomplexa had been named. They consisted of the gregarines (subclass Gregarinasida) (1624 named species and 231 named genera), the hemogregarines (family Haemogregarinidae) (399 species and 4 genera), the eimeriorins (order Eimeriorida) (1771 species and 43 genera), the hemospororids (order Haemospororida) (444 species and 9 genera), the piroplasmids (order Piroplasmorida) (173 species and 20 genera), and a few others (105 species and 32 genera). The first apicomplexan protozoon was seen by Antony van Leeuwenhoek; in 1674 he saw oocysts of Eimeria stiedai in the gall bladder of a rabbit. The first member of the phylum to be named (by Dufour in 1828) was Gregarina ovata in earwigs. During the quarter century 1826–1850, 41 species and 6 genera of Apicomplexa were named. These numbers increased progressively. In the quarter century 1951–1975, 1873 new species and 83 new genera were named. Data are given for the numbers of named species and genera of apicomplexan protozoa of each group known in 1850, 1875, 1900, 1925, 1950, 1975, and 1987.     

Gregarines of the Genus Lecudina (Protozoa,Apicomplexa) from Pacific Ocean Polychaetes*

The gregarine Lecudina longissima Hoshide, 1944 is described from the intestine of the marine polychaete Lumbrineris zonata (Johnson, 1901) from Santa Catalina Island, Calif. L. catalinensis n. sp. is described from the intestine of L. inflata Moore, 1911 from the same island. L. pherusae sp. n. and L. zimmeri sp. n. are described from the intestine of the marine polychaete Pherusa capulata (Moore, 1909) off Santa Catalina Island.     

Some corrections in Haemogregarine (Apicomplexa: Protozoa) nomenclature

The nomenclature of three genera in the family Haemogregarinidae (Haemogregarina, Karyolysus, and Hepatozoon) has been reviewed and the following new names are introduced to replace homonyms or for previously unnamed species: haemogregarina carlosi n. nom., in the erythrocytes of the lizard Lacerta ocellata; Haemogregarina tincae n. nom., in the stomach and intestine of the tench Tinca tinca; Hepatozoon insectivorae n. sp., in the leucocytes of the shrews Sorex araneus and Crocidura leucodon; Hepatozoon krampitzi n. sp., in the leucocytes of the vole Microtus oeconomus; Hepatozoon peromysci n. sp., in the leucocytes of the deermice Peromyscus boylii and P. truei gilberti; and Hepatozoon pallida (Pessoa et al., 1971) n. comb., in the erythrocytes of the snake Thamnodynastes pallidus nattereri.     

The Genera Barrouxia,Defretinella, and Goussia of the Coccidian Family Barrouxiidae (Protozoa,Apicomplexa)1

The apicomplexan family Barrouxiidae Léger, 1911 is reviewed and revised on the basis of present information. It includes the genera Barrouxia Schneider, 1885 with ten named species, Defretinella Henneré, 1966 with one named species, and Goussia Labbé, 1896 with 25 named species. The family is characterized by having bivalved sporocysts with a longitudinal suture line. Available information, admittedly spotty, is given for each species on oocyst, sporocyst and sporozoite structure, and on locus of sporulation. The following seven new combinations are made: Goussia flaviviridis (Setna & Bana, 1935) n. comb. in the gecko Hemidactylus flaviviridis; G. hyalina (Léger, 1898) n. comb. in an unidentified aquatic beetle; G. lacazei (Labbé, 1895) n. comb. in the centipedes Lithobius forficatus and L. martini; G. metchnikovi (Laveran, 1897) n. comb. in the gobies Gobio gobio and G. albipinnatus; G. schaudinniana (Pinto, 1928) n. comb. in the centipede Lithobius forficatus; G. stankovitchi (Pinto, 1928) n. comb. in the small bleak Alburnus alburnus, the bream Abramis brama, and the red roach Scardinius erythrophthalmus; Goussia sp. (Dogel' Akhmerov, 1959) nov. comb. in the freshwater fish Gnathopogon chankaensis.     

Some Corrections in Haemogregarine (Apicomplexa: Protozoa) Nomenclature1

The nomenclature of three genera in the family Haemogregarinidae (Haemogregarina, Karyolysus, and Hepatozoon) has been reviewed and the following new names are introduced to replace homonyms or for previously unnamed species: Haemogregarina carlosi n. nom., in the erythrocytes of the lizard Lacerta ocellata; Haemogregarina tincae n. nom., in the stomach and intestine of the tench Tinca tinca; Hepatozoon insectivorae n. sp., in the leucocytes of the shrews Sorex araneus and Crocidura leucodon; Hepatozoon krampitzi n. sp., in the leucocytes of the vole Microtus oeconomus; Hepatozoon peromysci n. sp., in the leucocytes of the deermice Peromyscus boylii and P. truei gilberti; and Hepatozoon pallida (Pessoa et al., 1971) n. comb., in the erythrocytes of the snake Thamnodynastes pallidus nattereri.     

Cross-Transmission of Eimeria spp. (Protozoa, Apicomplexa) of Rodents—A Review

A total of 169 cross-transmission attempts has been made with 44 (11.8%) of the 372 named species of Eimeria of rodents. Of these, 161 were rodent-to-rodent, 6 rodent-to-lagomorph, and 1 each rodent-to-carnivore and rodent-to-bird. None of the last three categories was successful. In the rodent-to-rodent combinations, 39 (80%) of the 49 attempts to transmit a coccidian species from one rodent species to another of the same genus were successful, and only 14 (12.5%) of the 112 attempts to transmit a coccidium to a rodent of a different genus were successful. Eight of the successful attempts were with E. chinchillae , which was the only truly euryxenous species of Eimeria in the group. Two successful attempts were between the closely related rodent genera Spermophilus and Cynomys , and two were both of E. separata from Rattus norvegicus to some genetic strains but not to others of Mus musculus. One attempt with E. vermiformis from Mus musculus to Rattus norvegicus required treatment of the rat with the immunosuppressant dexamethasone to succeed. More cross-transmission studies are needed to determine the host-spectra of the species of Eimeria and other coccidian genera, and to determine the roles of genetics and immunosuppression in their transmission.     

Named species and hosts of Sarcocystis (Protozoa: Apicomplexa: Sarcocystidae)

Summary A list is given of the present 93 species of the apicomplexan protozoan genus Sarcocystis together with their definitive and intermediate hosts (if known), synonyms, homonyms, lapsi calami, etc. The names of many species of this genus are poorly known, in doubt or controversial due to lack of access to some of the literature and to failure to accept the International Code of Zoological Nomenclature.The following taxonomic innovations are introduced: New species—S. nontenella for S. tenella Eble, 1961 [non] S. tenella (Railliet, 1886) from the buzzard Buteo buteo; S. scotti for Sarcocystis sp. from the housemouse completing its sexual development in the tawny owl, Strix aluco (see Tadros & Laarman, 1980); New combinations—S. ctenosauris for Cryptosporidium ctenosauris Duszynski, 1969 from the lizard Ctenosaura similis; S. lampropeltis for Cryptosporodium lampropeltis Anderson, Duszynski & Marquardt, 1968 from the king snake Lampropeltis c. calligaster; S. roudabushi for Isospora roudabushi Pellérdy, 1974 from the gopher snake Pituophis s. sayi; and S. tropicalis for Isospora tropicalis Mukherjea & Krassner, 1965 from the golden jackal Canis aureus.Supported in part by National Institute of Allergy and Infectious Diseases, National Institutes of Health, Bethesda, Maryland research grant AI15367.Supported in part by National Institute of Allergy and Infectious Diseases, National Institutes of Health, Bethesda, Maryland research grant AI15367.     

Redescription of Besnoitia bennetti (Protozoa: Apicomplexa) from the donkey (Equus asinus)

Besnoitia bennetti tissue cysts were found in four naturally-infected donkeys (Equus asinus) from the USA. Infectivity of its bradyzoites and tachyzoites to animals and cell culture was studied. The bradyzoites were not infectious to out-bred Swiss Webster mice, rabbits or gerbils. When fed tissue cysts, cats did not excrete oocysts. However, the parasite was infectious to interferon-gamma gene knock out mice. The parasite from tissues of two donkeys was grown successfully in bovine monocyte monolayers for the first time. Non-dividing, uninucleate tachyzoites were approximately 6 x 1.5 microm in size. Longitudinally-cut bradyzoites in tissue sections measured 8.7 x 1.9 microm. Ultrastructurally, tachyzoites and bradyzoites were similar to those in other Besnoitia species, and in particular to parasites described from cattle (Besnoitia besnoiti) and reindeer (Besnoitia tarandi), in that their bradyzoites lacked enigmatic bodies. Based on comparative analysis of three portions of nuclear ribosomal DNA (the small and large subunits and the first internal transcribed spacer) B. bennetti was found to be more closely related to the other congeners described from ungulates. The parasite was formally redescribed and specimens deposited in the US National Parasite Collections.     

Redescription of Besnoitia tarandi (Protozoa: Apicomplexa) from the reindeer (Rangifer tarandus)

Besnoitia tarandi tissue cysts were found in naturally-infected reindeer (Rangifer tarandus) from Finland. Infectivity of its tissue cysts, bradyzoites, and tachyzoites to animals and cell culture was studied. The bradyzoites and tissue cysts were not infectious to out-bred mice, rabbits or gerbils. When fed tissue cysts, neither cats nor dogs excreted oocysts. However, the parasite was lethal to interferon-gamma gene knock out mice irrespective of the route of inoculation. The parasite was grown successfully in African Green Monkey cells from tissues of two reindeer for the first time. Non-dividing, uninucleate tachyzoites from smears from cell cultures were 5.6 x 1.4 microm (4.5-7.4 x 1.0-1.9, n=50) in size. Longitudinally-cut bradyzoites in tissue sections measured 7.4 x 1.3 microm (6.5-7.8 x 1.0-1.6, n=30). Ultrastructurally, tachyzoites and bradyzoites were similar to those in other Besnoitia species, and in particular to parasites described from cattle (Besnoitia besnoiti) and equids (Besnoitia bennetti) in that their bradyzoites lacked enigmatic bodies. Based on comparative analysis of three portions of nuclear ribosomal DNA (the small and large subunits and the first internal transcribed spacer) B. tarandi was found to be more closely related to the other congeners described from ungulates. The parasite was formally redescribed and specimens deposited in the US National Parasite Collection.     

In vitro development of Neospora caninum (Protozoa: Apicomplexa) from dogs

The development of Neospora caninum isolated from naturally infected dogs was examined in mammalian cell cultures. Tachyzoites developed by endodyogeny when inoculated onto bovine monocyte or bovine cardiopulmonary artery endothelial cell cultures. Tachyzoites were 5.0 by 2.0 microns and had a posteriorly located nucleus. Cytopathogenic effects of parasite development consisted of the formation of holes in the cell monolayer associated with the rupture of infected host cells. Serial passage of tachyzoites was achieved by subinoculation of tachyzoites onto non-infected bovine monocyte cell cultures. It appears that N. caninum can be continuously grown in cell cultures.     

Sarcocystis neurona (Protozoa: Apicomplexa): description of oocysts, sporocysts, sporozoites, excystation, and early development

Equine protozoal myeloencephalitis is a major cause of neurological disease in horses from the Americas. Horses are considered accidental intermediate hosts. The structure of sporocysts of the causative agent, Sarcocystis neurona, has never been described. Sporocysts of S. neurona were obtained from the intestines of a laboratory-raised opossum fed skeletal muscles from a raccoon that had been fed sporocysts. Sporocysts were 11.3 by 8.2 microm and contained 4 sporozoites. The appearance of the sporocyst residuum was variable. The residuum of some sporocysts was composed of many dispersed granules, whereas some had granules mixed with larger globules. Excystation was by collapse of the sporocyst along plates. The sporocysts wall was composed of 3 layers: a thin electron-dense outer layer, a thin electron-lucent middle layer, and a thick electron-dense inner layer. The sporocyst wall was thickened at the junctions of the plates. Sporozoites were weakly motile and contained a centrally or posteriorly located nucleus. No retractile or crystalloid body was present, but lipidlike globules about 1 microm in diameter were usually present in the conoidal end of sporozoites. Sporozoites contained 2-4 electron-dense rhoptries and other organelles typical of coccidian zoites. Sporozoites entered host cells in culture and underwent schizogony within 3 days.     

Note on the Taxonomy of Frenkelia microti (Findlay & Middleton, 1934) (Apicomplexa: Sarcocystidae)

Based on biological, morphological and molecular data, species of Frenkelia Biocca, 1968 should be reclassified within the genus Sarcocystis Poche, 1913. This taxonomic change leads to the secondary homonymy of S. microti (Findlay & Middleton, 1934) n. comb. and S. microti Dubey, 1983. The recently suggested conspecificity and consequent synonymy of S. microti (Findlay & Middleton, 1934) and S. buteonis (Henry, 1932) is not really justifiable and thus S. microti (Findlay & Middleton, 1934) should be considered a valid species. S. jaypeedubeyi nom. nov. is, therefore, proposed as a nomen novum for S. microti Dubey, 1983 to alleviate the problem of this secondary homonymy.     

Infections in mice with tachyzoites and bradyzoites of Neospora caninum (Protozoa: Apicomplexa)

Tachyzoites of 2 isolates of Neospora caninum (NC-1 and NC-2) were inoculated subcutaneously (s.c.), intraperitoneally (i.p.), or orally into mice to compare the effects of route of inoculation on pathogenicity. Mice developed more severe disease, and disease occurred sooner when inoculated with the NC-1 isolate compared to the NC-2 isolate. Deaths occurred earlier in mice inoculated i.p. with either isolate. Mice inoculated orally or s.c. with tachyzoites responded similarly to infection. Tissue cysts of the NC-2 isolate produced infections in mice following oral or s.c. inoculation. Lesions seen in mice inoculated with tachyzoites or bradyzoites were primarily acute pneumonia, myositis, encephalitis, ganglioradiculoneuritis, and pancreatitis. In vitro studies demonstrated that tachyzoites of both isolates were killed by incubation in pepsin-HCl solution but not 1% trypsin solution. Bradyzoites of the NC-2 isolate were able to withstand treatment with pepsin-HCl solution.     

Goussia carpelli (Protozoa, Apicomplexa) in cyprinid fish of the Duero basin (NW Spain). Aspects of host-parasite relationships

Goussia carpelli was found parasitizing the intestine and gall bladder of Cyprinus carpio, Carassius carassius and Barbus barbus bocagei, and less frequently Leuciscus cephalus cabeda and Chondrostoma polylepis, from the Esla River. The coccidian is described and data on its prevalence, intensity and pathogenicity are given. In histological sections, the oocysts were detected in the lamina propria or submucosa. We did not observe a clear seasonal pattern nor an influence of the host sex on the infections. On the contrary, there was a tendency to a decrease in the prevalence with the age of the host.