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ISSUED SEPTEMBER, 1965 © 1965 by the Board of Trustees of the University of Illinois. Manufactured in the United States of America. Library of Congress Catalog Card No. 64-64686. CONTENTS INTRODUCTION TAXONOMY LIFE CYCLE GENERAL OOCYST MORPHOLOGY CRITERIA FOR SEPARATION OF SPECIES PATHOGENICITY SYSTEMATIC SECTION GENUS EIMERIA SCHNEIDER, 1875 GENUS ISOSPORA SCHNEIDER, 1881 GENUS DORISIELLA RAY, 1930 GENUS WENYONELLA HOARE, 1933 GENUS CARYOSPORA LEGER, 1904 GENUS TYZZERIA ALLEN, 1936 GENUS CRYPTOSPORIDIUM TYZZER, 1907 GENUS KLOSSIA SCHNEIDER, 1875 GENUS KLOSSIELLA SMITH AND JOHNSON, 1902 DISCUSSION SUMMARY LITERATURE CITED TABLES PLATES ADDENDUM INDEX a 3 P48 ‘ mt PeAee de ¥ sal eet ree Te ek Oe: veswey Ul sev de oe wp ASUTs vw ener INTRODUCTION Coccidia are among the commonest and most important protozoan parasites of vertebrates. Several hundred species belonging to over 30 genera have been described. Almost two hundred species belonging to 9 genera are known to occur in rodents. The literature on them is scat- tered. Some have been described well, others poorly. No critical analysis of the coccidia of the whole order has ever been made, and, as a result, anyone who wishes to study them is faced with a difficult and time-con- suming task. This was brought home to us when we began to study the coccidia of wild rodents in 1954, and the present monograph has grown out of that study. TAXONOMY Several schemes have been suggested for the classification of the coc- cidia (Wenyon, 1926; Reichenow, 1953; Grassé, 1953; Hall, 1953; Kudo, 1954; Cheissin, 1956; Hoare, 1933, 1957; Levine, 1961). The following classification, based on Levine (1961), includes the genera parasitic in rodents. The system of uniform endings for the names of higher taxa proposed by Levine (1959) is used. The ending, -asida, designates a class, -asina a subclass, -orida an order, and -orina a suborder. 2 THE COCCIDIAN PARASITES OF RODENTS Subphylum SPOROZOA (EUSPORA) Class TELOSPORASIDA Subclass GREGARINASINA Mature trophozoite extracellular, large. Subclass COCCIDIASINA Mature trophozoite intracellular (with a few exceptions), small. Order EUCOCCIDIORIDA Life cycle involves both sexual and asexual phases; schizogony present in the latter. Suborder ADELEORINA Macrogamete and microgametocyte associated in syzygy during differentiation. Few microgametes usually produced. Family ADELEIDAE Sporocysts formed in oocyst. In epithelium of gut and its appended organs. Chiefly in invertebrates. Genus KLOSSIA Schneider, 1875 Oocyst with many spherical sporocysts, each with three to ten sporozoites. Macrogametes not vermiform. Family KLOSSIELLIDAE Typical oocyst not formed; a number of sporocysts each with many sporozoites develop within a membrane which is perhaps laid down by the host cell. Two microgametes formed by microgametocyte. In kidney and other organs of host. Genus KLOSSIELLA Smith and Johnson, 1902 With the characters of the family. Suborder EIMERIORINA Macrogamete and microgametocyte develop independently. Microgametocyte typically produces many microgametes. No SAV EN Family CRYPTOSPORIDIIDAE Development within cuticular layer of host cell and not in cell proper. Oocyst without sporocysts. Genus CRYPTOSPORIDIUM Tyzzer, 1907 Oocyst with four naked sporozoites. Family EIMERIIDAE Development in host cell proper. Oocysts with none, one, two, four, or many sporocysts, each with one or more sporozoites. LIFE CYCLE 3 Genus EIMERIA Schneider, 1875 Oocyst with four sporocysts, each with two sporozoites. Genus ISOSPORA Schneider, 1881 Oocyst with two sporocysts, each with four sporozoites. Genus DORISIELLA Ray, 1930 Oocyst with two sporocysts, each with eight sporozoites. Genus CARYOSPORA Léger, 1904 Oocyst with one sporocyst containing eight sporozoites. Genus WENYONELLA Hoare, 1933 Oocyst with four sporocysts, each with four sporozoites. Genus TYZZERIA Allen, 1936 Oocyst with eight naked sporozoites. LIFE CYCLE The only genus of Adeleorina whose life cycle in rodents is known is Klossiella. Its life cycle is rather different from that of the Eimeriorina; it will be described in the systematic section below. The life cycles of the Eimeriorina are similar, and can be illustrated by that of Eimeria nieschulzi, which occurs in the small intestine of the rat (Fig. 1). (See Roudabush, 1937, Dieben, 1924, and Levine, 1957.) The oo- cysts are passed in the feces; at this time they contain a single cell, the sporont. In the presence of oxygen this cell elongates somewhat, throws off a polar granule by reduction division, rounds up again, and then di- vides into four sporoblasts, each of which then forms a sporocyst contain- ing two sporozoites. In E. nieschulzi the sporocysts contain a residual body while the oocysts do not; these bodies may be present or absent in other species. The sporulation process takes about three days in this species, but may take more or less time in other species. When a rat eats the sporulated oocysts, they break, releasing the sporozoites. These can be found in the lumen of the small intestine three to four hours after the rat has been infected. ‘They enter the cells of the intestinal epithelium and round up, becoming first-generation schizonts. These grow, and after about 36 hours form 20 to 36 first-generation merozoites by multiple fission. The merozoites break out of the epithelial cells, destroying them in the process. Each enters another cell, where it rounds up to form a second-generation schizont, grows, and produces 10 THE COCCIDIAN PARASITES OF RODENTS Fic. 1. Life cycle of Eimeria nieschulzi from Rattus norvegicus (from Levine ID), to 14 second-generation merozoites two days after infection. This process is repeated twice more, third- and fourth-generation merozoites being produced three and four days, respectively, after infection. The sporozoites are 10-12 « long and contain a nucleus at the center GENERAL OOCYST MORPHOLOGY 5 and an eosinophilic globule at either end. The merozoites do not have these globules nor do the sporozoites of all species of Eimeria. The first- generation merozoites are 7-10 u, the second-generation 13-16 u, the third-generation 17—22 u, and the fourth-generation ones 4—7 u long. The sexual phase of the life cycle follows this asexual phase. The fourth-generation merozoites enter new epithelial cells, round up, and form either microgametocytes or macrogametes. The microgametocytes produce a large number of biflagellated microgametes by multiple fission. The macrogametes simply grow. They contain a number of plastic gran- ules, which pass to their periphery, flatten out, and coalesce to form the oocyst wall after fertilization by a microgamete. The formation of this wall marks the transition of a fertilized macrogamete into an oocyst. The oocysts then break out of their host cells, enter the intestinal lumen, and pass out in the feces. The prepatent period, from the time of infection to the appearance of the first oocysts, is seven days. Oocysts continue to be discharged for four to five days, because the sporozoites do not all enter the host cells immediately, but may remain in the lumen of the intestine for four days before doing so. In the absence of reinfection, coccidial infections are self-limiting. Asexual reproduction does not continue indefinitely as it does, for ex- ample, in Plasmodium. In E. nieschulzi four generations of merozoites are produced; in other species there may be one, two, or three generations. After this, the life cycle enters its sexual phase, the oocysts are formed and eliminated from the body, and the infection is over. Reinfection may take place, but the host develops more or less immunity following the primary infection. GENERAL OOCYST MORPHOLOGY Since most coccidian species are differentiated on the basis of the morphology of their oocysts, the characters of the sporulated oocyst de- serve attention. These are shown in the drawing of a sporulated oocyst of Eimeria in Figure 2. Not all species have all the characters shown, and their presence or absence helps differentiate the species. The oocysts vary in shape. The oocyst wall is made up of one, two, or rarely three layers. Within it there may or may not be a lining membrane. A term which has been misused in descriptions of oocysts is “oval.” This term means “egg-shaped,” i.e., with one end broader than the other, 6 THE COCCIDIAN PARASITES OF RODENTS MICROPYLE CAP MIGROPYLE POLAR GRANULE STIEDA BODY SMALL REFRACTILE GLOBULE IN SPOROZOITE LARGE REFRACTILE GLOBULE IN SPOROZOITE SPOROCYST OOCYST RESIDUUM SPOROCYST RESIDUUM SPOROZOITE NUCLEUS SPOROZOITE INNER LAYER OF OOCYST WALL SPORULATED OUTER LAYER OF OOGYST WALL EIMERIA OOCYST Fic. 2. Sporulated oocyst of Eimeria (from Levine, 1961a). but it is often used when both ends of the oocyst are of equal width; the more proper term in this case is “‘ellipsoidal.’”” We have used the term “ovoid” in place of ‘oval’; this, too, means egg-shaped. However, because of the erroneous meaning often attributed to these terms, it must be recognized that oocysts which some authors have described as oval or ovoid may actually be ellipsoidal. Occasionally, too, some authors even use the term “egg-shaped” instead of “ellipsoidal”; for instance, Musaev and Veisov (196la) labeled the drawing of a subspherical oocyst of Eimeria gliris “oval’naya’” (i.e., oval) and an ellipsoidal oocyst “‘yait- sevidnaya”’ (i.e., egg-shaped). There may be a micropyle at one end which is covered by a micropylar cap in some species. Inside the oocyst are the sporocysts. In addition, a refractile polar granule and an oocyst residuum may be present. The oocyst residuum may be a compact body or may consist of a few to many, more or less scattered, granules or globules. The sporocysts also vary in shape. The sporocyst wall may vary in thickness and may have a knob (the Stieda body) at one end. The sporozoites are found inside the sporocysts. A sporocyst residuum may also be present as a compact body or consisting of a few to many, more or less scattered, granules. The sporozoites are usually elongate, with one end broader than the CRITERIA FOR SEPARATION OF SPECIES 7 other. The nucleus is either central or near one end. ‘There may be one or two, more or less refractile, globules in each sporozoite; if there are two, they are usually of unequal size. CRITERIA FOR SEPARATION OF SPECIES Both morphological and biological characteristics are used to separate the species of coccidia. There are differences in the morphological char- acters of both the endogenous and exogenous stages of the life cycle. However, since the endogenous stages of many species are unknown, the structure of the oocyst is most commonly used. As will be seen in the systematic section of this monograph, many descriptions of coccidian oocysts are sadly deficient. Too often nothing is said about the presence or absence of certain structures such as the Stieda body or sporocyst residuum, the shapes of the oocysts are not clearly de- scribed, those of the sporocysts are often not even mentioned, and the illustrations, if present, are so sketchy as to be almost useless. “The number of layers in the oocyst wall may not be mentioned or may be stated ambiguously. In this connection, use of the term “double-contoured,” should be discontinued. This phrase is extremely ambiguous. Some au- thors mean by it that the wall is composed of two layers, while others mean that there is a single layer whose inner and outer edges are heavily delineated. With some microscopes, spherical aberration is such that it is difficult to be sure how many layers are present because one or more extra lines appear to be present around the wall. Even if a microscope with an apochromatic oil immersion objective is used, the best way to be sure of the number of layers is to crush the oocyst. The structural details of coccidian oocysts are often so fine that use of an apochromatic objective is desirable in all studies of them. A second group of criteria used in differentiating coccidian species is the location of the endogenous stages in the host. Most species are found in the intestine, but some occur in other organs such as the liver and kidney. Within the intestine itself, different species are found in dif- ferent regions. One may be limited to the cecum, another to the duo- denum, another to the ileum, and still another may be found in more than one place. The location of the parasites within the host tissues or cells may differ 8 THE COCCIDIAN PARASITES OF RODENTS with different species even in the same region of the intestine. Some may be found in the epithelial cells at the tips of the villi, others only in the crypts of Lieberkithn, and still others in the submucosa of the villi. Within the host cell, some species may occur above the host cell nuclei (i.e., between the nucleus and the lumen of the intestine), others beneath them, and still others within them. Host specificity is a third criterion used in separating coccidian species. This varies wth the protozoan genus and to some extent wth the species. In general, the host range of Jsospora is relatively broad. Several members of the same host order may be infected by the same Jsospora species. For example, Isospora bigemina has been reported from the dog, cat, ferret, and mink. The host range of Eimeria is relatively narrow. A single species rarely infects more than one host genus unless the latter are closely related. Cross-immunity studies are also used in differentiating the coccidia of a particular host species. Infection of a host animal with one species of coccidium produces immunity against that species but not against other species which occur in the same host. The feeling is sometimes expressed that coccidia have so few structures that not many species can be distinguished morphologically. However, conservative calculation shows that at least 2,654,736 * morphologically different oocysts are possible in the genus Eimeria alone, to say nothing of differences in host, location in host, etc. (Levine, 1962). PATHOGENICITY While many species of coccidia are pathogenic, many others are not. Pathogenicity depends on a number of factors, some of which are prob- ably still unknown. Among those which might be mentioned are the number of host cells destroyed per infecting oocyst (which depends upon the number of merozoite generations and the number of merozoites * This figure is based on the assumption that the following numbers of pos- sibilities exist for the characters named: Oocyst shape, 4; oocyst size, 2; oocyst color, 3; number of layers in oocyst wall, 2; thickness of oocyst wall, 2; texture of oocyst wall (smooth, rough, striated, etc.), 3; oocyst polar granule, 2; oocyst residuum presence and type, 3; micropyle presence and type, 4; micropyle cap, 2; sporocyst size, 2; sporocyst shape, 4; Stieda body presence and shape, 3: sporocyst residuum presence, 2; sporozoite position (lengthwise, crosswise, etc.), 2; sporozoite refractile globule presence, 2. SYSTEMATIC SECTION 9 per generation) and the location of the parasites in the host tissues and within the host cells. The size of the infecting dose or doses, the degree of reinfection, and the degree of acquired or natural immunity of the host are also important. If disease is present, the signs are those of a diarrheal enteritis. There may or may not be blood in the feces, depending on the parasite species and severity of infection. Affected animals gain weight poorly, become weak and emaciated, or may even die, depending again on the parasite species and the size of the infecting dose of oocysts. Those animals which recover develop an immunity to the particular species which infected them. However, this is not an absolute immunity, and recovered adult animals are often continuously reinfected so that they carry light infections which do not harm them but which make them a source of infection for the young. In addition, under conditions of stress their immunity may be broken down and they may suffer from the disease again. SYSTEMATIC SECTION The genera of coccidia are taken up separately in this section. Within each protozoan genus, the species are arranged according to host genus. The rodent classifications of Ellerman (1940, 1941) and Simpson (1945) and the names employed by Ellerman and Morrison-Scott (1951) and Hall and Kelson (1959) are used. At the end of the section, the principal morphological characteristics of the oocysts of each species are tabulated. Included in this monograph are some of the results of a study of the coccidia of wild Illinois rodents trapped during the spring and summer of 1958 in the vicinity of Sullivan, Illinois. They were collected as part of a leptospirosis survey under the direction of Drs. Deam H. Ferris and Lyle E. Hanson of the University of Illinois College of Veterinary Medicine and Dr. Rexford D. Lord, Jr. of the Ilinois State Natural History Survey. Material from 16 deermice (Peromyscus maniculatus and P. leucopus), 4 voles (Microtus ochrogaster), 5 Franklin ground squirrels (Spermophilus franklinit), 11 Norway rats (Rattus norvegicus) and 42 house mice (Mus musculus) was examined. No coccidia were found in the voles and ground squirrels. The findings in the deermice have already been reported (Levine and Ivens, 1960). Those in the Norway rats and house mice are given below. 10 THE COCCIDIAN PARASITES OF RODENTS A study was also carried out on coccidia in fecal samples from 9 Rattus hawatiensis collected in August, 1958, in the vicinity of Honokaa on the island of Hawaii and sent to us by Mr. Henri P. Minette, Bureau of Laboratories, State Department of Health, Hilo, Hawaii. In studying the coccidia from []linois rodents, the contents of the colons and ceca were obtained at autopsy and placed in 2.5% potassium bi- chromate solution, mixed thoroughly, placed in a thin layer in a petri dish, and kept at room temperature for a week in order to sporulate. If they had not done so in this time, they were allowed to remain for another week or more. They were then stored in a refrigerator. They were examined microscopically after sugar flotation, using a Leitz Ortholux microscope equipped with apochromatic objectives. At the time of autopsy, segments of duodenum, jejunum, anterior ileum, posterior ileum and cecum were fixed in Zenker’s fluid. They were later sectioned and stained with hematoxylin and eosin by Dr. Marjorie B. Losch, and examined histologically for endogenous stages of the coccidia. The hosts were identified by Dr. Ferris or by Dr. Donald F. Hoffmeister, Department of Zoology, University of Illinois. The fecal samples from R. hawatiensis were mixed with 2.5% potassium bichromate before shipment to Urbana. Upon receipt, they were handled in the same way as the other material. GENUS EIMERIA SCHNEIDER, 1875 This genus is characterized by the presence of four sporocysts, each containing two sporozoites, in each oocyst. Host Suborder SCIUROMORPHA Host Superfamily SCIUROIDEA Host Family SCIURIDAE Host Subfamily SCIURINAE Host Tribe SCIURINI EIMERIA SCIURORUM GALLI-VALERIO, 1922 (Plate 1, Figs. 1-4; Plate 33, Figs. 270, 271) Eimeria sciurorum Galli-Valerio, 1922: 344-347; Yakimoff and Gousseff, 1935b: 740-741; Bornand, 1937: 509-514; Pellérdy, 1954a: 475-580; (?) Yakimoff, Sokoloff, and Rastegaieff, 1931: 487-490. GENUS Eimeria SCHNEIDER, 1875 1] Eimeria sciurt Yakimoft and Terwinsky, 1931: 56-59 (an invalid emendation, with no nomenclatural status). [non] Eimeria sciurorum: Mdller, 1923: 1-23 (see E. moelleri); Knipling and Becker, 1935: 417-418 (see E. kniplingi); Roudabush, 1937a: 107-108 (see E. parasciurorum); RySavy, 1954: 131-174 (see E. neosciurt); Bond and Bovee, 1958: 225-229 (?) (see Eimeria sp. Bond and Bovee, 1958). Description: Oocysts cylindroid, usually with parallel sides. Oocyst wall smooth, thin, colorless, probably composed of a single layer. Micro- pyle very small, seldom visible. Oocysts 2415 u according to Galli- Valerio (1922) or 26-29 x 13-20 u, with a mean of 28x16 u, according to Pellérdy (1954a). Mean length-width ratio 1.6 (Galli-Valerio) or 1.75 (Pellérdy). Unsporulated oocyst contains a round sporont 12 uw in diameter. Sporocysts oval, 10-14 x 6-8 u (Pellérdy). Pellérdy (1954a) stated that the sporocysts have a fine micropyle; perhaps by this term he meant Stieda body. Yakimoff and Gousseff (1935b) did not illustrate a Stieda body, but their drawing is quite diagrammatic. One or two re- fractile polar granules present. Oocyst residuum absent. Neither Galli- Valerio (1922) nor Pellérdy (1954a) made any statement about the sporocyst residuum. According to Yakimoff and Gousseff (1935b), it is represented by some scattered granules. Sporulation Time: Three or four days according to Pellérdy (1954a). Schizogony and Gametogony: The details of schizogony and gametog- ony are unknown. Pellérdy (1954a) describes some of the intracellular stages in a mixed infection with three other species of Eimeria, but it is impossible to say which stages were those of E. scitwrorum. All the parasites were found above the host cell nuclei. Prepatent Period: According to Pellérdy (1954a), the prepatent period is seven days. Type Host: Sciurus vulgaris var. alpinus (European alpine squirrel). Other Hosts: Sciurus vulgaris (squirrel); “Eichhornchen.” Location: Small intestine. Pellérdy (1954a) found oocysts beginning a few inches from the pylorus. Geographic Distribution: Europe (Switzerland, Italy, Hungary, USSR). Pathogenicity: Galli-Valerio considered this species an important cause of death in wild squirrels. Pellérdy (1954a) observed diarrhea and slight dysorexia on the fourth to sixth days following experimental infection with mixed coccidia of which EF. sciurorum was the most abundant. Bornand (1937) observed marked hyperemia of the small intestine in a heavily infected $. vulgaris which he examined. Cross-Transmission Studies: Galli-Valerio (1922) attempted unsuccess- fully to transmit this species from Sciuwrus vulgaris to the white rat. Prevalence: No information. 12 THE COCCIDIAN PARASITES OF RODENTS Remarks: The original description of this species was so incomplete that a number of different forms from several species of squirrels have been assigned to it by different authors. While their more complete de- scriptions are not incompatible with Galli-Valerio’s sketchy one, some of them are incompatible with each other. E, sciurorum was originally described by Galli-Valerio (1922) from the alpine squirrel, Sciwrus vulgaris var. alpinus, in Switzerland. He later (1930) listed Sciurus vulgaris var. noire as a host, but this varietal name is a vernacular and not a scientific one. Of the six coccidia described by other authors under the name Eimeria sclurorum, a critical study indicates that only two should be assigned to this species. These are the E. scitwrorum described by Pellérdy (1954a) from Sciurus vulgaris in Hungary, and probably E. sciurorum described by Yakimoff and Gousseff (1935b) from an “Ezchhornchen” (no scientific name given) in Russia. ‘The forms described under this name by Moller (1923) from Sciurus (syn., Neosciurus) carolinensis in a German zoological garden, by Yakimoff, Sokoloff, and Rastegaieff (1931) from S. vulgaris in Russia, by Knipling and Becker (1935) from S. niger rufiventer from Iowa, and by Roudabush (1937a) from the flying squirrel, Glaucomys volans, from Iowa, belong to different species. Galli-Valerio’s (1922) complete description was: “Eimeria sciurorum n. sp. Im Darme einer Sciurus vulgaris var. alpina aus St. Loup (Waadt) habe ich eine Exmeria vom 24 x 15 u gefunden, deren Gestalt mehr zylindroid als ovoid war. Protoplasma als Kugel in der Mitte der Oocyste, von 12 u. Mikropyle sehr klein. In Wasser gesetzt, haben die Oocysten 4 Sporen mit je 2 Sporozoiten ergeben. Die Gestalt dieser Eimeria war ganz verschieden von derjenigen der EF. stiedat.” The form described under the name £. sciturorum Galli-Valerio by Moller (1923) from Sciurus (syn., Neosciurus) carolinensis in a German zoological garden differs from this species in possessing a large micropyle (4-6 x 1-1.5 u) and in lacking an oocyst polar granule. The form described as E. sciturorum by Yakimoff, Sokoloff, and Rastegaieff (1931) from S. vulgaris in Russia differs from this species in having a large micropyle 7.2 u in diameter. However, it appears to agree with it in all other respects. Two of the oocysts in the rather crude drawings of Yakimoff, Sokoloff, and Rastegaieff (1931) are shown without micropyles, and the other two have a thickened wall at one end which might be supposed to represent a micropyle. Pending further study, it is considered best to list this form as a questionable EF. sciwrorum. The form described as E. sciwrorum by Knipling and Becker (1935) GENUS Eimeria SCHNEIDER, 1875 13 from S. niger rufiventer in Iowa differs from this species in lacking oocyst polar refractile granules and in having an oocyst residuum and a large sporocyst residuum. The form described as E. sciturorum by Roudabush (1937a) from Glaucomys volans in Iowa differs from this species in having a prominent sporocyst residual body and in lacking a micropyle. In addition, its host belongs to a different subfamily from that of Sciwrus; since most rodent Eimerias seem to be quite host-specific, this increases the probability that it is a different species. The form described by Bond and Bovee (1958) from Scrurus carolinen- sis under the name E. sciurorum (?) differs from E. sctuwrorum in having an oocyst residuum. A more complete description of this form is needed be- fore its taxonomic position can be established. EIMERIA BOTELHOI CARINI, 1932 Eimeria botelhot Carini, 1932: 80-82. Description: Although Carini (1932) described the oocysts as oval; they are actually illustrated as piriform in his drawing. Oocysts 36 x 28 u with a large micropyle at the tapering end. Oocyst wall 3 u thick, com- posed of three layers, of which the outermost is yellowish and rough. Sporocysts elongate ellipsoidal, 19 x 9 u, without a Stieda body. Sporocyst residuum small, granular, lying between the two elongate, comma-shaped sporozoites. Oocyst residuum and polar granules absent. Sporulation time: Several days in 1% chromic acid. Schizogony: Schizogony takes place in the mucosal cells of the villi. The young schizonts in the cells are round, and occur above the host cell nucleus. They grow to a diameter of 25-30 u and produce a variable number (generally 12-20) of merozoites. The merozoites are fusiform, 10-12 u long, 4 u wide, and have a nucleus in the center. Gametogony: Gametogony takes place in the mucosal cells of the villi. The gametocytes lie above or beside the host cell nuclei. Each micro- gametocyte produces hundreds of intensely staining microgametes 4—5 u long. The mature macrogametes have a central nucleus and vacuolar cytoplasm. After fertilization, the oocyst wall is laid down rapidly. Prepatent period: Unknown. Host: Sciurus (Guerlinguetus) ingrami (squirrel). Location: Small intestine. Geographic Distribution: Brazil. Pathogenicity: Unknown. The parasitized cells are destroyed. Cross-Transmission Studies: None. Prevalence: Unknown. 14 THE COCCIDIAN PARASITES OF RODENTS EIMERIA FRANCHINIT BRUNELLI, 1935 Eimeria franchini Brunelli, 1935: 354-366. Description: Oocysts piriform, with a prominent micropyle at the small end. Oocyst wall thick, rough, yellow, composed of two layers. Oocysts 24 x 15 u (Hardcastle, 1943, gave the dimensions erroneously as Wed: SO SETS) Sporulation Time: The oocysts failed to sporulate in 2.5% potassium bichromate after 24 days, although four sporoblasts formed in some. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Sciurus vulgaris (squirrel). Location: Feces. Geographic Distribution: Europe (Italy). Pathogenicity: Unknown. Prevalence: Unknown. Remarks: Although this species is incompletely described, it clearly belongs to the botelhoi series, but differs from all the other species in squirrels by its small size. EIMERIA LUISIERI (GALLI-VALERIO, 1935) REICHENOW, 1953 Jarrina luisieri Galli-Valerio, 1935: 643-644. Eimeria (Jarrina) luisieri (Galli-Valerio). Reichenow, 1953: 1-850. Description: Oocysts ovoid, with one end round and the other shaped like a bottleneck. Oocyst wall thick, rough, yellowish. Number of layers in oocyst wall not given. Micropyle very distinct. Oocysts 33x24 u. Sporont of unsporulated oocysts 18 u in diameter. Sporocysts almost spherical, 9 x 7.5 u. Sporozoites piriform. Oocyst residuum present. Sporocyst residuum and oocyst polar granule not mentioned. Sporulation time: ‘Twenty days at room temperature on moist filter paper in a moist chamber. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Sciurus vulgaris var. alpina (alpine squirrel). Location: Intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. This species was described from the fluid intestinal contents of a squirrel which was found dead. Prevalence: Unknown. EIMERIA MIRA PELLERDY, 1954 (Plate 33, Fig. 272) Eimeria piriformis Lubimov, 1934: 1-108 [fide Pellérdy, 1954a: 475-480] (a junior homonym of £. piriformis Kotlan and Pospesch, 1934). GENUS Eimeria SCHNEIDER, 1875 15 Eimeria mira Pellérdy, 1954a: 475-480. [non] Eimeria piriformis: Kotlan and Pospesch, 1934: 215-217; Marotel and Guilhon, 1941: 321-328. Description: Oocysts piriform, with a short bottleneck at the end of which is a large micropyle 4-6 u in diameter. Oocyst wall rough, brown, composed of two layers 3 u thick. Oocysts 30-40 x 19-27 u according to Pellérdy (1954a), or 35-37 x 24 u according to Lubimov (1934). Oocyst residuum and polar granule absent. Sporocysts cigar-shaped, pointed at both ends, 19 x 6 u, with a large amount of residual material. Sporozoites comma-shaped. Sporulation time: Five to seven days. Schizogony and Gametogony: The details of schizogony and gametog- ony are unknown. Pellérdy (1954a) described some of the intracellular stages in the ileum of a squirrel in a mixed infection with three other species of Eimeria, but it is impossible to say which stages were those of E. mira. All the parasites were found above the host cell nuclei. Prepatent Period: Ten to eleven days. Type Host: Sciurus vulgaris (squirrel). Location: Small intestine. Pellérdy (1954a) found oocysts only in the ileum of a squirrel killed about 11 days after experimental infection. Geographic Distribution: Hungary, USSR. Pathogenicity: Unknown. Cross-Transmission Studies: Pellérdy (1954a) was unable to infect three ground squirrels (Spermophilus citellus) or three dormice (Glis glis) with E. mira from Sciurus vulgaris. Prevalence: Unknown. Remarks: This species was first described by Lubimov (1934) from Sciurus vulgaris under the name Eimeria piriformis. Kotlan (1951), how- ever, showed that this name was a homonym of Eimeria piriformis Kotlan and Pospesch, 1934 from the rabbit. Since he thought the squirrel species might be a synonym of Eimeria eubeckeri Hall and Knipling, 1935 from Franklin’s ground squirrel, Spermophilus franklinii, Kotlan (1951) did not assign it a new name. Later, however, Pellérdy (1954a) showed that it differed from E. eubeckeri in the shape of its micropyle, and was unable to infect the European ground squirrel, Spermophilus citellus, with oocysts from Sciurus vulgaris. He therefore gave this form the name Eimeria mira. The only differences between this species and FE. botelhoi Carini, 1932 aside from the host and geographic locality are that FE. mira has narrower sporocysts pointed at both ends and a larger amount of sporocyst residual material. However, Pellérdy did not illustrate his sporulated oocysts, nor did he refer to E. botelhoi or to Carini’s paper, so it is possible that future research may show that these two forms belong to a single species. 16 THE COCCIDIAN PARASITES OF RODENTS EIMERIA ANDREWSI YAKIMOFF AND GOUSSEFF, 1935 (Plate 33, Figs. 273 and 275) Eimeria andrewsi Yakimoft and Gousseff, 1935b: 740-741; Pellérdy, 1954a: 475-480. Description: Oocysts oval, somewhat pointed at both ends, or sub- spherical to ellipsoidal. Oocyst wall smooth, composed of a single layer. Oval oocysts 19.8-22.5 x 14.4-16.2 u (mean, 20.9 x 15.2 u); subspherical oocysts 18.0-20.1 x 15.3-16.2 uw (mean, 18.5x15.4 u). Length-width ratio of oval oocysts, 1.3—1.5 (mean, 1.4); length-width ratio of subspherical oocysts, 1.18—-1.25 (mean, 1.20). The above dimensions are from Yakimoft and Gousseff (1935b). According to Pellérdy (1954a), the oocysts measure 19-25 x 14-16 u. Oocyst polar granule present. Oocyst residuum absent. Sporocysts oval, 7.2 x 3.6 u, without Stieda body or residuum. Sporulation Time: Three days at room temperature, according to Pellérdy (1954a). Schizogony and Gametogony: The details of schizogony and gametog- ony are unknown. Pellérdy (1954a) described some of the intracellular stages in a mixed infection with three other species of Eimeria, but it is impossible to say which stages were those of FE. andrewsi. All of the parasites were found above the host cell nuclei. Prepatent Period: According to Pellérdy (1954a), the prepatent period is six days. Type Host: “Eichhornchen” (presumably Sczurus sp.). Other Hosts: Sciurus vulgaris (squirrel). Location: Throughout small intestine. Geographic Distribution: Europe (White Russia, Hungary, probably England). Pathogenicity: Unknown. Prevalence: Unknown. Remarks: ‘The unsporulated coccidium described and illustrated by Sheather (1923) from a squirrel in England probably belonged to this species. It was ellipsoidal, had a smooth, thin wall without a micropyle, and measured 21-25 x 12-16 u. EIMERIA SILVANA PELLERDY, 1954 (Plate 33, Fig. 276) Eimeria silvana Pellérdy, 1954a: 475-480. Description: Odocysts ellipsoidal or subspherical, 15-18 x 12-15 u. Oocyst wall smooth, pale; number of layers not stated. Micropyle absent. Entire unsporulated oocysts filled by sporont. Sporocysts elongate. Oocyst residuum absent. No information given on oocyst polar granule, sporocyst residuum, or Stieda body. GENUS Eimeria SCHNEIDER, 1875 17 Sporulation Time: One to two days. Schizogony and Gametogony: Unknown. Pellérdy (1954a) described some of the intracellular stages in a mixed infection with three other species of Eimeria, but is impossible to say which stages were those of E. stlvana. All the parasites were found above the host cell nuclei. Prepatent Period: Seven days. Type Host: Sciurus vulgaris (squirrel). Location: Small intestine. Geographic Distribution: Europe (Hungary). Pathogenicity: Unknown. Prevalence: Unknown. EIMERIA SERBICA POP-CENITCH AND BORDJOCHKI, 1957 (Plate 1, Figs. 5 and 6) Eimeria serbica Pop-Cenitch and Bordjochki, 1957: 73-75. Eimeria serbca [sic]: Ibid. Description: Oocysts described as oval and illustrated as ellipsoidal, yellowish rose, 21-35 x 12-25 u. Oocyst wall 0.6—0.9 u thick, illustrated as composed of a single layer. Micropyle absent. Oocyst residuum absent. Oocyst illustrated without polar granule. Sporocysts illustrated as el- lipsoidal, without Stieda body. Sporocyst residuum present, 3.5 uw in diameter. Sporulation Time: At least 12 days at 25C and 45 days at 5C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Serbian squirrel (Sciurus vulgaris ?). Location: Unknown. Oocysts found in feces. Geographic Distribution: Yugoslavia. Pathogenicity: Unknown. Prevalence: Unknown. Remarks: Although the description of this species is incomplete, and although Pop-Cenitch and Bordjochki (1957) were apparently unaware of any other squirrel coccidia except Eimeria sciurorum, their species appears to be different from the others reported from squirrels, except E. parasciurorum Bond and Bovee, 1957 (p. 45). However, the fact that the host of the latter, Glaucomys volans, belongs to a different subfamily from Sciurus vulgaris makes the probability of cross-transmission doubt- ful. This species is therefore retained pending a more detailed description and cross-transmission study. EIMERIA MOELLERI N. SP. Eimeria sciurorum Galli-Valerio. Moller, 1923: 1-23. [non] Eimeria scturorum Galli-Valerio, 1922: 345-347. 18 THE COCCIDIAN PARASITES OF RODENTS Description: Oocysts ellipsoidal, sometimes cylindrical, 22-28 x 14-18 u. Oocyst wall smooth, composed of two layers. Micropyle 4—6 u wide and I-1.5 uw high. Sporocysts 10-14 x 6-8 u, with a small micropyle (Stieda body?) at one end. Sporocyst residuum present. Oocyst polar granule and residuum absent. Sporozoites dumbbell-shaped with one end somewhat pointed, 8-14 x 4-5 u. Sporulation Time: Three days. Schizogony and Gametogony: Moller (1923) described these processes in an experimentally infected Scrurus vulgaris. The schizonts are more or less round and about 18 u in diameter. They enlarge the host cell, pushing its nucleus to one side. Each schizont forms 12 or more merozoites measuring 6-7 x 1-2 u. The microgametocytes are round and 16-20 u in diameter. They form a large number of slender, comma-shaped microgametes. The macrogametes are round and 16-20 u in diameter, with a star- or thornapple-shaped nucleus. Prepatent Period: Thirteen days. Type Host: Sciurus (Neosciurus) carolinensis (gray squirrel). Location: Small intestine, especially posterior part of jejunum. Para- sitic in the tips of the villi. Geographic Distribution: Europe (described from an American squirrel in the Berlin Zoological Garden). Pathogenicity: Nonpathogenic. Cross-Transmission Studies: Moller (1923) transmitted this coccidium experimentally to a European domestic squirrel (presumably Sciurus vulgaris). Prevalence: Unknown. Remarks: Although Moller (1923) called this form E. sciwrorum and was followed by a number of other authors, it clearly belongs to a different species. It differs from E. sciwrorum especially in possessing a large micropyle 4-6 u in diameter rather than a generally inapparent one. In addition, it has no oocyst polar granule, while E. sciwrorum does. It is also somewhat smaller. The other species of squirrel Eimeria with a prominent micropyle are E. botelhoi, E. franchini, E. luisieri, E. petauristae, and E. mira, but all of these have a thick, rough, yellowish to brownish wall, whereas that of Mdller’s species is thin, smooth, and apparently colorless. The remaining named species of squirrel Eimeria, all of which have thin, smooth walls, all lack a micropyle. In addition, MOller’s species differs from E. andrewsi in lacking an oocyst polar ganule, in having a sporocyst residuum, and in having larger sporocysts. It is much larger than EF. glaucomydis. Finally, it differs from £. silvana in being much larger and in having a sporocyst residuum. GENUS Eimeria SCHNEIDER, 1875 19 We are therefore assigning the name Ezmeria moelleri n. sp. to the species described under the name Eimeria sciturorum by Moller (1923) from Sciurus (Neosciurus) carolinensis. EIMERIA NEOSCIURI PRASAD, 1960 (Plate 1, Figs. 7-14; Plate 2, Figs. 15 and 16) Eimeria neosciuri Prasad, 1960: 135-139. Eimeria sciurorum Galli-Valerio. RySavy, 1954: 131-174. [non] Eimeria neosciuri: Webster, 1960: 139-146 (see E. ascotensis). [non] Eimeria sciurorum Galli-Valerio, 1922: 344-347. Description: Oocysts ellipsoidal, 22-28 x 14-18 u, with a length-width ratio of 1.55-1.58. Oocyst wall smooth, composed of two layers. Outer layer colorless or pale yellowish; inner layer dark brown. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, with protruding Stieda body, 11-13 x 5-7 u. Sporocyst residuum present. Sporozoites described as sickleshaped, 6 < 2-3 u, illustrated as comma-shaped with a clear globule at the large end. Sporulation Time: Thirty-six to forty-eight hours at 22C. Schizogony and Gametogony: According to Prasad (1960), the schizonts are oval, 10-11 x 4-4.5 u, and contain 10-13 nuclei. The merozoites are sausage-shaped, 5—5.5 x 1 u, and have a central nucleus. The microgametocytes are asymmetrical, ellipsoidal, or cylindrical, and measure 19-20 x 12 u. They produce a large number of microgametes, which are arranged haphazardly around a central residual mass. The macrogametes are ovoid or cylindroid, measure 5-17 x 5-12 u, and have a central, vesicular nucleus and a series of plastic granules around their periphery. The number of schizont generations and the length of the patent period are unknown. Prasad (1960) followed the wave of oocyst produc- tion in one squirrel after a relapse. It was highest from the sixth to the tenth days and then decreased gradually; no more oocysts were passed after about two weeks. Prepatent Period: Unknown. Type Host: Sciurus (Neosciurus) carolinensis (gray squirrel). Other Host: Sciurus vulgaris (squirrel). Location; Small intestine. According to Prasad (1960), all stages occur beneath the host cell nucleus in the epithelial cells of the villi of the upper part of the ileum. Geographic Distribution: Europe (England, Czechoslovakia). Pathogenicity: Prasad (1960) saw no macroscopic lesions, but remarked that the affected part of the ileum appeared slightly thinner than the noninfected part. Heavily infected squirrels showed no signs of illness, and their feces and appetites were normal. 20 THE COCCIDIAN PARASITES OF RODENTS Cross-Transmission Studies: None. Prevalence: Prasad (1960) found this species in all of four squirrels which he examined in England. Remarks: Rysavy (1954) reported finding “Eimeria sciurorum” in Sciurus vulgaris in Czechoslovakia. However, the oocysts which he il- lustrated had a prominent Stieda body and resembled those of E. neosciurt more than they did those of E. sciturorum. They were ellipsoidal to ovoid and measured 24-32 x 14-19 u, with a mean of 28 x 18 u and a mean length-width ratio of 1.67. Their oocyst wall was thin and colorless and reported to be composed of a single layer. There was no distinct micropyle. An oocyst polar granule was present (but rarely) and an oocyst residuum was absent. The sporocysts were ovoid and 8-12 x 6—7 u. A sporocyst residuum was present. The sporulation time was 72 hours, and the parasites were reported to occur in the small intestine. EIMERIA ASCOTENSIS N. SP. (Plate 2, Figs. 17-24) Eimeria neosciurt Prasad. Webster, 1960: 139-146. [non] Eimeria neosciuri Prasad, 1960: 135-139. Description: Oocysts (illustrated as ellipsoidal) ovoid, 14-31 x 10-20 u, with a mean of 24 x 15 u. Oocyst wall apparently smooth, composed of three layers: outer layer pale yellow, 0.75 u thick; middle layer brown, 1.0 uw thick; and inner layer pinkish orange, 0.6 uw thick (but the oocyst wall in the photomicrograph appears thinner than this and composed of only one layer). True micropyle absent, its place taken by an operculum 6-9 u wide; in addition, at the end of the oocyst opposite the operculum is a rather indistinct, wavy line 5-7 u long in the two inner layers of the wall, which Webster (1960) called a terminal cap, but which looks like a second operculum. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, with a prominent Stieda body, 9-10 x 6-7 u. Sporocyst residuum present. Sporozoites piriform, 9 x 3 u, with a clear globule at the large end. Sporulation Time: Ninety-five per cent sporulate in 76 hours at 20, 25, and 30C, in about 140 hours at 15C, and in about 192 hours at 10C. After 24 hours, about sixty-five per cent of the oocysts were sporulated at 30C, about forty-five per cent at 25C, and about five per cent at 20C. The oocysts died in 24 hours at 40C. Schizogony and Gametogony: According to Webster (1960), the schizonts are 8-17 uw in diameter and contain 6 to 11 crescentic or banana- shaped merozoites measuring 4.5-6 x 1.5 u. The microgametes measure 2.5-3 x 0.6-0.9 u. The macrogametes are GENUS Eimeria SCHNEIDER, 1875 21 spherical and were said by Webster to be 6-8 u in diameter; these were probably immature. Prepatent Period: Unknown. Type Host: Sciurus (Neosciurus) carolinensis (gray squirrel). Location: Small intestine. According to Webster (1960), all the intra- cellular stages occur in the epithelial cells distal to the host cell nucleus. Geographic Distribution: Europe (England). Pathogenicity: According to Webster (1960), infected squirrels showed no signs of disease and their feces were normal. However, in all the squirrels he examined the duodenum was rather inflamed and _ the jejunum frequently had swollen lymph nodes and small hemorrhages. In one squirrel he noted that the gametogony phase of the infection was associated with a large collection of lymphocytes between the sub- mucosa and the mucosa which appeared as an opaque white patch on the ileum. Cross-Transmission Studies: None. Prevalence: Webster (1960) found this species in all of six squirrels which he examined from Silwood Park, Sunninghill, Ascot, Berkshire, England. Remarks: Webster (1960) considered this species to be Eimeria neosciuri, but his description differs from Prasad’s (1960) descrip- tion of FE. neosciuri in several important respects. If both descriptions are correct, then the two forms must be different species. The oocysts of Webster’s species had an operculum and a “terminal cap,” while those of Prasad’s species had neither. Webster said that his form had a three- layered oocyst wall whereas Prasad said that his had a two-layered wall. (It is possible that the outermost layer of Webster’s oocysts may have been an optical illusion, since the operculum and terminal cap did not ex- tend through it and since we have seen this optical illusion in many oocysts of various species which we ourselves have studied; therefore, we do not place much emphasis on this character.) The sporocysts of Webster’s form are smaller than those of Prasad’s. In addition, the location of the endogenous stages in the host cells differed. Prasad found these stages beneath the host cell nuclei, while Webster found them above the nuclei. Since Webster’s description differs from those of all other species of Eimeria from squirrels, we are naming his form Eimeria ascotensis n. sp.; it is named after Ascot, where the infected squirrels were found. E. ascotensis differs from all other squirrel Eimerias in having an operculum and “terminal cap.” It differs further from E, botelhoi, E. no ho THE COCCIDIAN PARASITES OF RODENTS franchiniu, E. luisiert, E. petauristae, and E. mira in lacking a thick, rough, yellowish to brownish wall and a true micropyle. It differs further from E. sciwrorum in oocyst shape and in having a prominent Stieda body. It differs from E. andrewsi and E. serbica in having a prominent Stieda body. It differs further from E. moelleri in lacking a true micropyle and in having an oocyst polar granule. It differs further from E. kniplingi in oocyst shape, in having an oocyst polar granule, and in lacking an oocyst residuum. It differs further from E. silvana in oocyst size. EIMERIA KNIPLINGI N. SP. (Plate 3, Figs. 25-27) Eimeria sciurorum Galli-Valerio. Knipling and Becker, 1935: 417-418. [non] Eimeria sciurorum Galli-Valerio, 1922: 344-347. Description: Oocysts cylindrical with rounded ends, 15.5-34.0 x 10.1- 19.0 u, with a mean of 24.2 14.2 u; length-width ratio, 1.35-2.15, with a mean of 1.70. Oocyst wall smooth, tinted pinkish to orange, composed of a single layer. No micropyle visible, but oocyst wall appeared to be slightly thinner at one end. Sporont almost fills unsporulated oocyst. Sporocysts ellipsoidal, 13.1 x 6.7 u, with a prominent Stieda body, a large sporocyst residual body, and additional residual material. Oocyst polar granule absent. Oocyst residuum present, small. Sporulation Time: Fifty-seven to seventy hours in potassium bichro- mate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Sciurus niger rufiventer (fox squirrel). Location: Cecum and large intestine; a few in small intestine. Geographic Distribution: United States (Iowa). Pathogenicity: Unknown. Prevalence: Unknown. Remarks: This coccidium was described from a single fox squirrel in Ames, Iowa. It was assigned somewhat hesitantly to E. sciurorum, but in view of the additional information now available on the latter species, it is clear that Knipling and Becker’s form does not belong to it. It differs in lacking oocyst polar granules and in having an oocyst residuum, a large sporocyst residuum, and a prominent Stieda body. In addition, its habitat is primarily the large intestine, while that of EF. sciurorum is the small intestine. This species differs from E. moelleri in lacking a micropyle and in having an oocyst residuum and a prominent Stieda body. In addition, it occurs primarily in the large intestine, while E. moelleri occurs primarily in the jeyunum. It differs from E. andrewsi in shape, in lacking an oocyst polar granule, GENUS Eimeria SCHNEIDER, 1875 23 in having both oocyst and sporocyst residua, and in having a Stieda body. In addition, the sporocysts are much larger. It is larger than FE. stlvana and differs from it in shape. It differs also in having oocyst and sporocyst residua and a Stieda body. It is larger than E. glaucomydis and differs from it in shape. It differs also in having an oocyst residuum and a Stieda body. It differs from E. serbica and E. parasciurorum in having an oocyst residuum and a sporocyst Stieda body. It differs from FE. neosciuri in the location of its endogenous stages, in having an oocyst residuum, and in lacking an oocyst polar granule. It differs from EF. botelhoi, E. franchiniu, E. luisiert, E. petauristae, and E. mira in lacking a micropyle and having a smooth, pale, thin wall, while all these have thick, rough, yellowish to brownish walls. There are other differential characters, but it is not necessary to give them. We are therefore assigning the name, Eimeria kniplingi n. sp. to the species described under the name Eimeria sciturorum by Knipling and Becker (1935) from Sciurus niger rufiventer. EIMERIA SP. HENRY, 1932 Eimeria sp. Henry, 1932a: 279-290. Description: Oocyst ovoid, 22.4—-32.0 x 16.0-19.2 u, with a mean of 28.8 x 19.2 u. Oocyst wall smooth. Micropyle present. Sporulated oocysts not described. Sporulation Time: About two days. Host: Sciurus griseus griseus (gray squirrel). Location: Oocysts found in cecal contents. Geographic Distribution: United States (California). Prevalence: Unknown. Remarks: This form was described from a single gray squirrel. Henry (1932) stated that it might be different from E. scturorum, but that there was not sufficient evidence to differentiate it. She thought that it was similar to the unsporulated oocyst figured from a squirrel in England by Sheather (1923); this latter form was probably E. andrewsi. Further in- formation is needed before Henry’s form can be assigned to any species. EIMERIA SP. BRUNELLI, 1935 Eimeria sp. Brunelli, 1935: 357-360. Description: Oocysts ovoid, with a smooth, pearl gray double wall, 17.5 x 9.75 u. A granular residual body is present at one of the poles, but it is not clear from Brunelli’s description whether this is an oocyst or a sporocyst residuum. Sporulation Time: Four to six days in 2.5% potassium bichromate. Schizogony and Gametogony: Unknown. 24 THE COCCIDIAN PARASITES OF RODENTS Prepatent Period: Unknown. Type Host: Sciurus vulgaris (squirrel). Location: Feces. Geographic Distribution: Europe (Italy). Pathogenicity: Unknown. Prevalence: Unknown. Remarks: Further morphological information is needed before this form can be assigned to any species. EIMERIA SP. BOND AND BOVEE, 1958 (Plate 3, Fig. 28) Eimeria sciurorum (?) Galli-Valerio. Bond and Bovee, 1958: 225-229. Description: Oocysts illustrated as ellipsoidal, with a single-layered wall. Oocysts 27 x 17 u. Small micropyle present. Oocysts illustrated with- out polar body but with oocyst residuum. Sporocysts illustrated as elongate kidney-shaped, without Stieda body or sporocyst residuum. Schizogony and Gametogony: Unknown. Type Host: Sciurus carolinensis (grey squirrel). Location: Feces. Geographic Distribution: North America (Florida). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. Remarks: Bond and Bovee (1958) made a drawing of this form but did not give a complete description. They mentioned its size and the presence of a small micropyle and said that it was almost identical in size and shape to E. sciurorum as described by Galli-Valerio (1922). How- ever, it differs from this species in having an oocyst residuum and in lacking a polar granule. Bond and Bovee did not refer to Pellérdy’s (1954a) work on squirrel coccidia. Their form does not agree with the description of any named species, but a more complete description is needed before its taxonomic position can be established. Host Suborder SCIUROMORPHA Host Superfamily SCIUROIDEA Host Family SCIURIDAE Host Subfamily SCIURINAE Host Tribe TAMIASCIURINI EIMERIA TAMIASCIURI LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 3, Fig. 29) Eimeria tamiasciuri Levine, Ivens, and Kruidenier, 1957: 80-88; Bullock, 1959: 39-40; Dorney, 1962: 258-261. GENUS Eimeria SCHNEIDER, 1875 25 Description: Oocysts elongate ellipsoidal, occasionally somewhat ovoid, rarely with one side slightly concave and the other convex. Oocyst wall smooth, colorless, composed of a single layer 1.5 u thick. Micropyle absent. Fifty sporulated oocysts measured 26-40 x 16-21 u, with a mean of 32.5 x 19.3 u. Their length-width ratios ranged from 1.4—2.0, with a mean of 1.69. Sporocysts elongate ovoid, about 168 u, very thin-walled at the sides, and broadly thickened at one end, and tapered to a blunt point at the other, thus forming a Stieda body. A single oocyst polar granule present, often lying among the sporocysts and difficult to see. Oocyst residuum absent. Sporozoites elongate and bent, embedded in a mass of residual material which fills the sporocyst. Sometimes this residual material forms a large body in the center of the sporocyst. Four hundred oocysts measured by Dorney (1962) from four Wisconsin red squirrels were 19-39 x 10-22 u, with a mean of 26x15 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Tamiasciurus hudsonicus (red or spruce squirrel). Location: Intestinal contents. Dorney (1962) found oocysts throughout the entire small intestine, with an especially large number in the “posterior quarter (ileum).” Geographic Distribution: United States (Arizona, New Hampshire, Wisconsin). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described from a single spruce squirrel from the south rim of Grand Canyon. Bullock (1959) found it in 2 road-killed T. hudsonicus loquax in New Hampshire. Dorney (1962) found it in 98 per cent of 50 T. hudsonicus in Wisconsin. EIMERIA TODDI DORNEY, 1962 (Plate 3, Fig. 30) Eimeria toddi Dorney, 1962: 258-261. Description: Oocysts “oval” (illustrated as ellipsoidal). Fifty-one sporulated oocysts measured 36—45 x 27-36 u, with a mean of 40 x 32 u; their length-width ratios averaged 1.25. Oocyst wall composed of two layers, the outer one rough, deep yellow, about 2.4 u thick; the inner one colorless, 0.6 4 thick. Micropyle absent. Zero to five (mean, 1.5) oocyst polar granules present. Oocyst residuum absent. Sporocysts ovoid, with prominent Stieda body and sporocyst residuum. ‘Twenty sporocysts measured 16-20 x 7-13 u, with a mean of 19x 11 u. Sporozoites each with one terminal and one central clear globules. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. 26 THE COCCIDIAN PARASITES OF RODENTS Prepatent Period: Not more than 12 days. Type Host: Tamiasciurus hudsonicus (red squirrel). Location: Intestinal contents. Geographic Distribution: North America (Wisconsin). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Dorney (1962) found this species in 6 per cent of 36 T. hudsonicus in northeastern Wisconsin. Host Suborder SCIUROMORPHA Host Superfamily SCIUROIDEA Host Family SCIURIDAE Host Subfamily SCIURINAE Host ‘Tribe XERINI EIMERIA GARNHAMI McMILLAN, 1958 (Plate 3, Figs. 31-35; Plate 4, Figs. 36 and 37) Eimeria garnhami McMillan, 1958: 20-23. Description: Oocysts spherical to subspherical. Oocyst wall smooth, pink, 0.7 u thick, composed of an outer thin layer and an internal thick one. Oocysts 14-19 x 13-18 u, with a mean of 15.5 14.7 u. Micropyle absent. McMillan (1958) stated, ‘““There are numerous refractile bodies, 0.5 u in diameter, arranged at the periphery in two layers. These peripheral bodies are probably left over after the wall of the oocyst is laid down from the refractile bodies of the macrogametocyte.” In his figures, however, these “refractile bodies” are shown within the cytoplasm of the sporont and not in the sporulated oocyst. They are therefore actually plastic granules rather than polar bodies, and a true polar body is absent. Oocyst residuum absent. Sporocysts spherical, smooth, 6.5 u in diameter. Sporocyst residuum present. Sporozoites 5.8 x 1.8 u, curved, slightly tapered at one end, with a central nucleus. Sporulation Time: Fourteen to sixteen days at 28-40C. No sporulation occurred at room temperature (21—22C). Schizogony: Intracellular stages toward apex of host cell. Host cell nucleus seldom distorted. Mature schizonts with six to eight elongate, pointed merozoites measuring 7.5 x 1.2 u. Gametogony: Only early microgametocytes were seen. They measured up to 6.58.0 u, but their multiple, rounded nuclei had not divided into microgametes. The macrogametes (‘“‘macrogametocytes’’) were spherical, 10.2 uw in diameter, with many spherical bodies 2.0 u in diameter around the periphery. Prepatent Period: Unknown. GENUS Eimeria SCHNEIDER, 1875 27 Sporogony: Not described. Type Host: Xerus (Euxerus) erythropus (African ground squirrel). Location: Epithelial cells of villi of at least the proximal part of the small intestine. Geographic Distribution: Africa (Katsina Province, Northern Nigeria). Pathogenicity: McMillan (1958) observed neither signs of illness nor macroscopic or microscopic pathologic changes in the intestines of affected animals. Cross-Transmission Studies: None. Host Suborder SCIUROMORPHA Host Superfamily SCIUROIDEA Host Family SCIURIDAE Host Subfamily SCIURINAE Host Tribe MARMOTINI EIMERIA MARMOTAE GALLI-VALERIO, 1923 Eimeria marmotae Galli-Valerio, 1923: 120-125; Bornand, 1937: 509-514. Description: Oocysts ovoid with somewhat rounded ends, 51x42 u, with a very distinct micropyle and containing a sporont 33 u in diameter. Sporulated oocysts not described. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Marmota (syn., Arctomys) marmota (marmot). Location: Intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described by Galli-Valerio (1923) from a single marmot. Bornand (1937) found it in two out of six M. marmota in Switzerland. Remarks: Despite Galli-Valerio’s sketchy description, this species can be differentiated from all others described from the genus Marmota by its large size. Bornand (1937) gave no further description. EIMERIA ARCTOMYSI GALLI-VALERIO, 1931* Eimeria arctomysi Galli-Valerio, 1931: 98-106. Description: Odocysts cylindroid, 24x20 u. Micropyle clearly visible and protruding a little. Sporont 16 y in diameter. Sporulated oocysts not * The genitive of mys is myis, not myst. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 28 THE COCCIDIAN PARASITES OF RODENTS described except to indicate that they contained four sporocysts each with two sporozoites. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Marmota (syn., Arctomys) marmota (marmot). Location: Intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA MONACIS FISH, 1930 (Plate 4, Figs. 38 and 39; Plate 33, Fig. 278) Eimeria monacis Fish, 1930: 98-100; Iwanoft-Gobzem, 1934: 149-151. Eimeria dura Crouch and Becker, 1931: 127-131. Description: Oocysts spherical to subspherical, 17-23 x 15-21 u, with a mean of 20.0x 18.3 u and a mean length-width ratio of 1.09, according to Fish (1930). Crouch and Becker (1931) reported that the oocysts were 14-20 u in diameter. Oocyst wall smooth, colorless, composed of a single layer. Micropyle absent. Sporont fills unsporulated oocyst. Sporu- lated oocysts without polar granule but with rather small oocyst residuum. Sporocysts ovoid, with Stieda body and sporocyst residuum. The form found by Iwanoff-Gobzem (1934) in M. bobak was ovoid, 17-21 x 13-18 u, with a mean of 17.8x 15.2 u. An oocyst residuum was present. No other information was given. Sporulation Time: Sixty to sixty-four hours at room temperature in 2% potassium dichromate solution according to Crouch and Becker (1931). Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Marmota monax (woodchuck). Other Hosts: Marmota bobak (marmot). Location: Intestine (found in feces and cecal contents). Geographic Distribution: United States (Washington, D.C., Iowa); USSR (Kazakhstan). Pathogenicity: Unknown. Hill (1952) reported that coccidiosis caused the death of a woodchuck (M. monax) in the London zoo, but did not describe the organism. Cross-Transmission Studies: None. Prevalence: Iwanoft-Gobzem (1934) reported this species from one out of two M. bobak in Kazakhstan. Remarks: Crouch and Becker (1931) remarked that they had described this form in manuscript under the name E. dura, but that when Fish’s (1930) description of E. monacis appeared they recognized that their form belonged to this species. GENUS Eimeria SCHNEIDER, 1875 29 EIMERIA OS CROUCH AND BECKER, 1931 (Plate 34, Fig. 279) Eimeria os Crouch and Becker, 1931: 127-131. Description: Oocysts ovoid, 20-26 x 18-22 u. Oocyst wall smooth, color- less, composed of a single layer. Micropyle distinct; sometimes the mem- brane lining the oocyst wall protrudes through the micropyle, forming a bulblike swelling. Sporulated oocysts without either polar granule or oocyst residuum. Sporocysts ovoid to ellipsoidal, with small Stieda body and sporocyst residuum, 9-13 by 5-8 u. Sporulation Time: Ninety to one hundred and five hours in 2% potassium bichromate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Marmota monax (woodchuck). Location: Intestine (found in feces and cecal contents). Geographic Distribution: United States (Iowa). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA PERFOROIDES CROUCH AND BECKER, 1931 (Plate 34, Fig. 280) Eimeria perforoides Crouch and Becker, 1931: 127-131. Description: Oocysts ellipsoidal, 17-24 x 15-20 u. Oocyst wall smooth, colorless, composed of a single layer. Micropyle absent. Sporont consid- erably contracted from oocyst wall even in fresh material. Sporulated oocysts without polar granule but with a small oocyst residuum. Sporo- cysts with Stieda body and sporocyst residuum, approximately 10x 4 u. Sporulation Time: Seventy hours in 2% potassium bichromate solu- tion at room temperature. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Marmota monax (woodchuck). Location: Intestine (found in feces and cecal contents). Geographic Distribution: United States (Iowa). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA CYNOMYSIS ANDREWS, 1928 * (Plate 4, Fig. 40) Eimeria cynomysis Andrews, 1928: 193-194. Eimeria cynomysi [sic] Ibid. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 30 THE COCCIDIAN PARASITES OF RODENTS Description: Oocysts broadly ellipsoidal, 33-37 x 28-32 u, with a mean of 35.4-30.0 u and a mean length-width ratio of 1.18. Oocyst wall 1.5—2.5 u thick, composed of two layers, the outer one transparent, with a fibrous appearance and a very irregular outer surface, the inner layer faint orange-yellow. Micropyle 5-6 u in diameter. Oocyst residuum absent. No oocyst polar granule mentioned or illustrated. Sporocysts seed-shaped, 13-17 x 8-12 u, with an inconspicuous Stieda body. Sporocyst residuum coarsely granular. Sporozoites attenuated reniform, blunt at both ends, but with one end a trifle larger than the other. Sporulation Time: Three to four days in moist feces at room tem- perature. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cynomys ludovicianus (prairie dog). Location: Intestine (found in feces). Geographic Distribution: United States (in prairie dogs from an animal supply house in Ohio). Pathogenicity: Unknown. Cross-Transmission Studies: Andrews (1927) reported that Eimeria sp. (presumably E. cynomysis) from the prairie dog could not be trans- mitted to the cat. Prevalence: This species was found in both of the two animals ex- amined. EIMERIA CITELLI KARTCHNER AND BECKER, 1930 (Plate 4, Figs. 41-44; Plate 5, Figs. 46-48; Plate 35, Fig. 291) Eimeria citelli Kartchner and Becker, 1930: 90-94; Sassuchin and Rauschen- bach, 1932: 646-650; Zasukhin and Tiflov, 1932: 129-132; Zasukhin and Tiflov, 1933: 437-442; Yakimoff and Sokoloff, 1935 (1934): 331-334; Zolotarev, 1938: 658-661; Pellérdy and Babos, 1953: 167-172; RySavy, 1957: 331-336; Svanbaev, 1962: 23-39. Description: Oocysts subspherical, ellipsoidal, or ovoid, 15-33 x 14-19 u, with a mean of 18.8 15.8 u. The oocysts described by Sassuchin and Rauschenbach (1932) from Spermophilus (syn., Citellus) pygmaeus meas- ured 17.4—26.1 x 14.5-19.6 u, with a mean of 21.1x17.2 u; the ovoid forms described by Yakimoff and Sokoloff (1934) from S. pygmaeus measured 17.0—21.6 x 14.4-18 u, with a mean of 19.2 16.8 u; the round forms described by Yakimoff and Sokoloff (1935) from the same host measured 14.4—23.4 u, with a mean of 16.9 u; the ovoid forms described by Zolotareff (1938) from S$. pygmaeus measured 15-24 x 13-21 u, with a mean of 19.2 x 16.2 u; the round forms described by him from the same host measured 15-21 u, with a mean of 17.1 u; those described by Pellérdy GENUS Eimeria SCHNEIDER, 1875 31 and Babos (1953) from S. citellus measured 15.0—22.9 x 14.0-20.3 u; and the nonspherical ones described by RySavy (1957) from S. citellus meas- ured 17-23 x 13-19 u, with a mean of 20x16 u, while the spherical ones were 13-19 uw in diameter, with a mean of 17 u. Oocyst wall smooth, colorless, composed of an intermediate thick layer and thin endo- and ecto-membranes. Micropyle absent. Freshly sporulated oocysts contain an oocyst residuum which becomes inconspicuous within three or four days. Oocyst polar granule absent. Sporocysts 5-9 4-7 u, with a small Stieda body. Sporocyst residuum present. Sporozoites 5—7.5 x 1.8-3.3 u, with a large refractile globule at the broader, more rounded end. Sporulation Time: Three days in 4% potassium bichromate. Schizogony: Kartchner and Becker (1930) observed schizogony in S. tridecemlineatus, but did not describe it. In S. citellus, Pellérdy and Babos (1953) reported that the schizonts lie above the host cell nuclei. The mature schizonts are round or subspherical, 10-13 u in diameter, and contain six to eight falciform merozoites with a central nucleus. The free merozoites measure 4—6 x 1.3—1.5 u. Gametogony: Although Kartchner and Becker (1930) did not describe gametogony in S. tridecemlineatus, they stated that the gametocytes usually lie above the host cell nucleus, but may be either below it or to the side. According to Pellérdy and Babos (1953), the mature macro- gametes from S. citellus are oval, 16x 11 u, and have a row of eosinophilic granules around their periphery which later coalesce. The microgame- tocytes are not numerous. They are round or elliptical, and measure 129 u. Each microgametocyte produces 50 to 70 microgametes measur- ing 2-3 x 0.3 u; no flagella were seen. Prepatent Period: Four to five days. Type Host: Spermophilus (syn., Citellus) tridecemlineatus (thirteen- striped ground squirrel). Other Hosts: Spermophilus (syn., Citellus) pygmaeus, S. citellus, S. maximus (ground squirrels). Location: Kartchner and Becker (1930) stated that the parasitic stages occur in the mucosa of the cecum of S. tridecemlineatus. Pellérdy and Babos (1953) on the other hand, stated that while gametogony occurs sporadically in epithelial cells of the cecum and colon, the small intestine, and particularly the jejunum and ileum, is the most common site of infection in S. citellus. Geographic Distribution: North America (Iowa), USSR (Kazakhstan, RSFSR, Saratoff, Krim, Daghestan), Europe (Hungary, Czechoslovakia). Pathogenicity: Although Kartchner and Becker (1930) observed no un- favorable effects in natural and heavy experimental infections of S. tridecemlineatus, Pellérdy and Babos (1953) observed catarrhal enteritis 32 THE COCCIDIAN PARASITES OF RODENTS of the small intestine and in some cases of the large intestine in experi- mentally infected S. citellus, and noted diarrhea and inappetance in one animal and death in another. Zolotareff (1938), too, considered in- testinal coccidiosis to be a cause of death in S. pygmaeus. Cross-Transmission Studies: Kartchner and Becker (1930) were unable to infect white rats and mice with E. citelli from S. tridecemlineatus. Prevalence: Kartchner and Becker (1930) found this species in 20.5 per cent of 78 S. tridecemlineatus in Iowa. Zasukhin and Tiflov (1932, 1933) reported that it was common in S. pygmaeus in southeastern RSFSR. Svanbaev (1962) found this species in 5 per cent of 43 S. maximus in southern Kazakhstan, USSR. EIMERIA BEECHEYI HENRY, 1932 (Plate 33, Fig. 277) Eimeria beecheyi Henry, 1932a: 279-290. [non] Eimeria beecheyi: Tanabe and Okinami, 1940: 126-134 (see E. astatict). Description: Oocysts ovoid, 16-22 x 10-13 u, with a mean of 19.2% 16.0 u. Oocyst wall smooth, colorless, composed of a single layer 1 u thick. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocyst residuum absent. Sporulation Time: Four to five days in 2% potassium bichromate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) beecheyi (ground squirrel). Location: Intestine. The oocysts were found in the contents of the cecum and large intestine. Geographic Distribution: United States (California). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described from two ground squirrels from Lake County and the vicinity of Nevada City, California. Remarks: Tanabe and Okinami (1940) described an Eimeria from Eutamias asiaticus in Japan under the name Eimeria beecheyt, but its oocysts contained sporocyst residua, while those of E. beecheyi do not, and the difference in host also suggests that the two species are different. We are therefore assigning it the new name, Eimeria asiatici (see below). EIMERIA BILAMELLATA HENRY, 1932 (Plate 5, Fig. 49; Plate 34, Figs. 285 and 286; Plate 35, Fig. 287) Eimeria bilamellata Henry, 1932a: 279-290; Pellérdy and Babos, 1953: 167-172. Eimeria eubeckeri Hall and Knipling, 1935: 128-129; RySavy, 1957: 331-336. Description: Since this coccidium has been described from three differ- ent hosts, separate descriptions are given for each host. In the form GENUS Eimeria SCHNEIDER, 1875 33 described by Henry (1932a) from Spermophilus lateralis, the oocysts are ovoid, 25-36 x 22-26 u, with a mean of 32.0 25.6 u. Oocyst wall com- posed of two layers: the outer one brown, thick, and rough; and the inner one clear, thin, and smooth. Micropyle present; oocyst wall thinner at the micropylar end. Sporulation time, less than ten days. Sporulated oocysts without oocyst residuum. Sporocysts 16.0 x 9.6 u, with two sporo- zoites and a large amount of residual material. In the form described by Pellérdy and Babos (1953) from S. citellus, the oocysts measure 30-41 x 22-28 u, and have an oocyst polar granule but no oocyst residuum. Their sporulation time was seven to eight days. Pellérdy and Babos (1953) gave no further description of the oocysts, since they agreed with Henry’s description. In the form described by RySavy (1957) from S. citellus, the oocysts were said to measure 34.2 [sic]—38 x 25-27 u, with a mean of 33.6 [sic] x 26 u; the majority were 30x27 u. They were broadly ovoid with a rough, strong, brownish yellow wall, with a prominent micropyle 6 u in diam- eter, without an oocyst residuum, with ovoid sporocysts 13x 9.5 u, and with a coarsely granular sporocyst residuum. Their sporulation time was 92 hours at 24C. In the form described by Hall and Knipling (1935) from S. franklinit, the oocysts measure 28—40 x 21-32 u, with a mean of 33.7 x 23.8 u. They are ovoid, with a thick wall composed of two layers, the outer one very rough and brown, and the inner one smooth and pinkish. A pronounced micropyle is present. The sporulation time in 4% potassium bichromate was 10 to 11 days. Oocyst polar granule and oocyst residuum absent. Sporocyst Stieda body and a large sporocyst residuum present. Sporulation Time: See above. Schizogony and Gametogony: Unknown. Prepatent Period: Pellérdy and Babos (1953) found this to be eight days in S. citellus. Type Host: Spermophilus (syn., Citellus) lateralis (syn., Callosper- mophilus chrysodeirus) (golden-mantled ground squirrel). Other Hosts: Spermophilus (syn., Citellus) citellus (ground squirrel); S. franklini (Franklin ground squirrel). Location: This species was found by Henry (1932a) in the feces of S. lateralis. It was found by Pellérdy and Babos (1953) in the small intestine, and particularly in the jejunum and ileum of S. citellus, and by Hall and Knipling (1935) in the cecum and large intestine of S. franklinii. Geographic Distribution: United States (California, Iowa), Europe (Hungary, Czechoslovakia). Pathogenicity: Unknown. Cross-Transmission Studies: None. 34 THE COCCIDIAN PARASITES OF RODENTS Remarks: This species was found by Henry (1932a) in two S. lateralis from Placer County, California, and by Hall and Knipling (1935) from a single S. franklinit in Iowa. The latter authors named their form Eimeria eubeckeri, but on the basis of the published descriptions and illustrations, it is morphologically indistinguishable from E. bilamellata and has approximately the same sporulation time. Hall and Knipling made no mention of Henry’s paper, and were presumably unaware of it. Cross-transmission studies would be necessary to prove whether the forms from the three hosts are identical, but in the absence of any other differentiating characters, it is preferable to assign them to the same species. EIMERIA CALLOSPERMOPHILI HENRY, 1932 (Plate 6, Fig. 54; Plate 34, Figs. 281 and 282) Eimeria callospermophili Henry, 1932a: 279-290; Levine, Ivens, and Krui- denier, 1958: 291-298. Eimeria callosphermophili [sic]: Svanbaev, 1962: 23-39 (in Spermophilus maximus). [non] Eimeria callosphermophili [sic]: Ibid. (in Meriones tamariscinus) (see E. assaensis). Description: According to Henry’s (1932a) original description of this species from the type host, the oocysts are subspherical, 16.0—22.4 x 16.0— 22.4 uw, with a mean of 19.2 16.0 u. Oocyst wall slightly rough and yellowish, probably composed of two layers. Micropyle absent, sporont almost filling the unsporulated oocyst. Oocyst polar granule present. Oocyst residuum large, homogeneous, and clear or granular, 3-5 u in diameter. Sporocysts almost round, pointed at one end, 10.2x8.5 u, containing two sporozoites and a few residual granules. In the form described by Levine, Ivens, and Kruidenier (1958) from Spermophilus (syn., Citellus) spilosoma spilosoma the oocysts are spherical to subspherical, 15-27 x 14-25 u, with a mean of 20.1 x 19.0 u; length-width ratios range from 1.0-1.1, with a mean of 1.06. Oocyst wall colorless to pale yellowish, slightly rough and pitted, composed of a single layer about 1.1 u thick. Micropyle absent. Sporocysts lemon-shaped, about 9X7 u, with Stieda body. Oocyst polar granule present. Oocyst residuum composed of several large, homogeneous bodies. Sporocyst residuum absent or com- posed of 1 to 15 or more round granules. Sporozoites often at the ends of the sporocysts but sometimes lying lengthwise in them. Levine, Ivens, and Kruidenier (1958) also found oocysts which con- tained only two sporocysts each with two sporozoites and thus resembled Cyclospora, but they resembled E. callospermophili in all other char- acteristics. In addition, abnormal oocysts of other types were present; these included oocysts with three normal sporocysts, with two normal and one giant sporocyst, or with one, two, three, or four imperfectly GENUS Eimeria SCHNEIDER, 1875 3 Or developed sporocysts. They therefore concluded that the Cyclospora-like forms were actually abnormal E. callospermophili. Sporulation Time: Six to seven days in 2% potassium dichromate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) lateralis (syn., Callosper- mophilus chrysodeirus) (ground squirrel). Other Hosts: Spermophilus (syn., Citellus) spilosoma spilosoma (spotted ground squirrel), Spermophilus (syn., Citellus) maximus (suslik). Location: Intestine. Henry (1932a) found the oocyst in the intestinal contents, particularly those of the cecum. Levine, Ivens, and Kruidenier (1958) found the oocysts in the intestinal contents. Geographic Distribution: United States (California) (type host); Mex- ico (Aguascalientes) (S. spilosoma); USSR (Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaey (1962) found this species in 14 per cent of 43 S. maximus in southern Kazakhstan. Remarks: This species was first described from S. lateralis in Placer County, California. The form described by Levine, Ivens, and Kruidenier (1958) from S. spilosoma in Mexico differs from it somewhat. Its oocyst wall is composed of a single layer, while that of the form from S. lateralis was said to be “probably” composed of two layers, although they could not be separated. There is obviously an error in the figures for oocyst width given by Henry (1932a), but whether it is the range or the mean which is wrong cannot be determined. The sporocysts of the form from S. lateralis are somewhat larger than those from S. spilosoma. Finally, Henry’s (1932a) description makes no mention of the Stieda body or of the position of the sporozoites in the sporocysts, and these cannot be determined from her photomicrographs. However, in the absence of a more detailed description of the form from the original host species, these differences do not appear to warrant the creation of a new species for the form from S. spilosoma. Svanbaev (1962) described a coccidium under the name E. callospher- mophili [sic] from Meriones tamariscinus. It does not belong to this species, and we have assigned it the name E. assaensis (p. 90). EIMERIA VOLGENSIS SASSUCHIN AND RAUSCHENBACH, 1932 (Plate 5, Fig. 50; Plate 34, Figs. 283 and 284) Eimeria volgensis Sassuchin and Rauschenbach, 1932: 646-650; Zolotareff, 1938: 658-661; Zasukhin and Tiflov, 1932: 129-132; Zasukhin and Tiflov, 1933: 437- aoe [non] Eimeria volgensis: Svanbaev, 1956: 180-191 (see E. kazakhstanensis). 36 THE COCCIDIAN PARASITES OF RODENTS Description: Oocyst ovoid, sometimes with a sharply pointed end, 23-32 x 17-28 u, with a mean of 27.2 x 21.9 u. The oocysts measured by Zolotareff (1938) were 18-24 x 15-20 u, with a mean of 21.2x16.4 u. Oocyst wall smooth, clear, colorless or greenish, composed of two layers, 1.2 u thick, becoming thinner at the micropylar end. Micropyle present, sometimes sharply pointed. If the micropyle is flat, it is 4-6 u in diameter. Sassuchin and Rauschenbach (1932) saw no sporulated oocysts; the follow- ing description is taken from Zolotareff (1938). Oocyst residuum absent. Sporocysts pear-shaped, 9-12 x 4-6 u. Sporocyst residuum present. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) pygmaeus (ground squirrel). Location: Intestine. Geographic Distribution: USSR (western Kazakhstan, RSFSR, Dagh- estan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Sassuchin and Rauschenbach (1932) found this species in 1 out of 242 rodents (most of which were S. pygmaeus) examined in western Kazakhstan. Zasukhin and Tiflov (1932, 1933) reported that this species was common in S. pygmaeus in southeastern RSFSR. Remarks: Even though the form described by Zolotareff (1938) was smaller than that originally described by Sassuchin and Rauschenbach (1932), Zolotareff considered it to be the same species. Since the original authors saw no sporulated oocysts, their morphology cannot be compared, but the sharply pointed micropylar end of both forms and their occur- rence in the same host and the same country tend to confirm Zolotareft’s view. Svanbaev (1956) described an Eimeria from the mole lemming, Ellobius talpinus, under the name E. volgensis. However, for the rea- sons given below, we do not consider it to belong to this species and have assigned it the name EF. kazakhstanensis (p. 86). EIMERIA BECKERI YAKIMOFF AND SOKOLOFF, 1935 (Plate 36, Fig. 292) Eimeria beckeri Yakimoff and Sokoloff, 1935: 331-334; Zolotareff, 1938: 658- 661; Svanbaev, 1956: 180-191 (in Spermophilus pygmaeus); Svanbaev, 1962: 23-29. Eimeria ussuriensis Yakimoft and Sprinholtz-Schmidt, 1939: 117-123. [non] Eimeria beckeri: Svanbaev, 1956: 180-191 (in Ellobius talpinus) (see E. talpint). Description: Oocyst usually ovoid, sometimes spherical. Ovoid forms 18-24 x 15-23 u, round forms 16-24 u in diameter. The oocysts found by GENUS Eimeria SCHNEIDER, 1875 37 Svanbaev (1956) in Spermophilus pygmaeus in Kazakhstan measured 29-98 x 19-24 u, with a mean of 25.5x 21.0 u. Oocyst length-width ratio 1.1-1.4, with a mean of 1.20. Oocyst wall yellow, smooth, composed of two layers. Micropyle absent. Sporont almost fills unsporulated oocysts. Sporulation time unknown. Oocyst polar granule and oocyst residuum absent. Sporocysts ovoid, seldom pinion 6—l2>c— 7 u, without Stieda body, but with a sporocyst residuum about 4 u in diameter. The oocysts of the form described from Spermophilus eversmanni by Yakimoff and Sprinholtz-Schmidt (1939) under the name Eimeria ussuri- ensis were mostly ovoid and oval, but some were spherical or subspherical. The ovoid oocysts measured 22—24 x 18-20 u, with a mean of 22.6 19.5 u; the oval oocysts measured 21—28 x 18-23 u, with a mean of 23.4 x 20.0 u; the subspherical oocysts measured 21-25 x 19-22 u, with a mean of 22.9 x 20.3 u; the spherical oocysts measured 18-25 u in diameter, with a mean of 22.1 uy. Oocyst wall yellow, composed of a single layer 1.2 u thick. Micropyle absent. Oocyst polar granule and oocyst residuum absent. The authors stated that they did not know whether there was a sporocyst residuum. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) pygmaeus (ground squirrel). Other Hosts: Spermophilus eversmanni (ground squirrel), Spermophi- lus (syn., Citellus) maximus (suslik). Location: Intestine (oocysts found in feces). Geographic Distribution: USSR (Crimea; Daghestan; Kazakhstan; Ussuri; Far Eastern Siberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: According to Zolotareft (1938), this species is widely dis- tributed in ground squirrels in Daghestan. Svanbaev (1956) found it in 58 per cent of 19 S. pygmaeus in western Kazakhstan. Svanbaev (1962) found it in 9 per cent of 43 S. maximus in southern Kazakhstan. Remarks: In their description of E. ussuriensis, Yakimoff and Sprin- holtz-Schmidt (1939) stated that it resembled E. beckeri but could be distinguished from it by its larger dimensions, which reached 32.4 x 27.0 u. However, this was the size of a single oocyst, and it was so exceptional that it was not even included in the series of ranges they listed. These ranges were similar to that of FE. beckeri. Thus, there is no valid reason for considering this form a separate species on the basis of present evidence. Svanbaev (1956) described an Eimeria from the mole lemming, Ellobius talpinus, under the name E. beckert. However, for the reasons given 38 THE COCCIDIAN PARASITES OF RODENTS below, we do not consider it to belong to this species and have assigned it the name E. talpini (p. 86). EIMERIA FRANKLINIT HALL AND KNIPLING, 1935 (Plate 5, Fig. 51) Eimeria franklinii Hall and Knipling, 1935: 128-129. Description: Oocysts subspherical to ovoid, 19-24 13-18 u, with a mean of 20.6 15.2 u. Oocyst wall smooth, colorless, transparent, com- posed of two layers. Micropyle absent. Sporulated oocysts with two polar granules and an oocyst residuum. Oocyst residuum decreases in size with age. Sporocysts ellipsoidal to subovoid, 11.56 u, with Stieda body. Sporocyst residuum present. Sporulation Time: Three to four days at room temperature in potas- sium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) franklini (Franklin ground squirrel). Location: Intestine. Oocysts were found in the contents of the small intestine, cecum, and colon. Geographic Distribution: United States (Iowa). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described from a single ground squirrel at Ames, Iowa. EIMERIA LATERALIS LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 6, Fig. 58) Eimeria lateralis Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ellipsoidal to somewhat ovoid. Fifty-nine oocysts measured 28-40 x 24-31 u, with a mean of 35.1 x 26.8 u; their length- width ratios ranged from 1.0-1.5, with a mean of 1.25. Oocyst wall 2 u thick, yellowish brown, rough, pitted like a thimble, and composed of a single layer. A thin membrane lines the oocyst wall, making its inner margin appear heavier than its outer one. Micropyle absent. No com- pletely sporulated oocysts were observed. The great majority contained a single cell, and most of those which had developed had done so ab- normally and contained two, three, or sometimes four rounded or irregularly shaped bodies. Some oocysts, however, contained four sporo- cysts measuring about 16x 10 u, a granular oocyst residuum, and one or more polar granules. Sporocysts, with prominent Stieda body, were filled GENUS Eimeria SCHNEIDER, 1875 39 with round residual granules and a few clear spherules. No definite sporozoites were observed. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) lateralis (mantled ground squirrel). Location: Intestinal contents. Geographic Distribution: United States (Grand Canyon National Park, Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA HOFFMEISTERI LEVINE, IVENS, AND KRUIDENIER, 1958 (Plate 5, Figs. 52 and 53) Eimeria hoffmeisteri Levine, Ivens, and Kruidenier, 1958: 291-298; Ivens, Kruidenier, and Levine, 1959: 53-57. Description: Oocysts subspherical. Oocyst wall colorless to pale yellow- ish, smooth, composed of a single layer about 1.0 u thick. Micropyle absent. Seven sporulated oocysts measured 15-20 x 13-18 u, with a mean of 17.7 16.1 u. Their length-width ratios ranged from 1.0-1.2, with a mean of 1.10. Sporocysts elongate ovoid, about 11.5 6.0 u; Stieda body absent or inconspicuous. Oocyst polar granule present. Oocyst residuum absent or composed of a few homogeneous bodies. Sporocyst residuum composed of a large amount of finely granular material. Sporozoites either at the ends of the sporocysts or lying somewhat longitudinally. Twenty-nine sporulated oocysts measured by Ivens, Kruidenier, and Levine (1959) from a Spermophilus spilosoma trom Chihuahua, Mexico, measured 17-23 x 16-19 u, with a mean of 19.9 17.2 u. Their length- width ratios ranged from 1.0-1.3, with a mean of 1.14. Very slight differ- ences were noted in the appearance of the sporocyst residual material and Stieda body between these oocysts and those in the original host. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) s. spilosoma (spotted ground squirrel). Location: Intestinal contents. Geographic Distribution: Mexico. The original host was trapped near Rincon de Romos, Aguascalientes. The same species was later found by Ivens, Kruidenier, and Levine (1959) in a S. spilosoma from Barenda, Chihuahua. 40 THE COCCIDIAN PARASITES OF RODENTS Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA SUSLIKI N. SP. Eimeria ussuriensis Yakimoft and Sprinholtz-Schmidt. Svanbaev, 1962: 23-29. [non] Eimeria ussuriensis Yakimoff and Sprinholtz-Schmidt, 1939: 117-123 (see E. beckert). Description: Oocyst ellipsoidal or elongate ellipsoidal, 29-35 x 23-26 u, with a mean of 32x25 u. Oocyst length-width ratio 1.2—-1.4, with a mean of 1.3. Oocyst wall smooth, 1.0-1.2 u thick, greenish, lilac, or yellow-brown. Micropyle absent. Oocyst polar granule and residuum absent. Sporocysts ellipsoidal or short ellipsoidal, 9-13 x 7-10 u, witha mean of 109 u. Sporocyst residuum absent. Sporozoites comma-shaped, 6-9 x 3-5 u, with a mean of 8x4 u. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) maximus (suslik). Location: Oocysts found in feces. Geographic Distribution: USSR (southern Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962) found this species in 5 per cent of 43 S. maximus in southern Kazakhstan. Remarks: Svanbaev (1962) called this form Ezmeria ussuriensis. How- ever, E. ussuriensis is a synonym of E. beckeri, and the form that Svanbaev found resembles neither the original description of E. becker, nor Yakimoff and Sprinholtz-Schmidt’s (1939) description of E. us- suriensis, nor the description of any other species of Eimeria heretofore reported from Spermophilus. Hence, it is necessary to give it a new name, and we are therefore calling it Eimeria susliki n. sp. EIMERIA SP. LEVINE, IVENS, AND KRUIDENIER, 1957 Eimeria sp. Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts broadly ovoid to ellipsoid. A single oocyst meas- ured 1917 u, with a length-width ratio of 1.1. Oocyst wall smooth, pale tan, 0.9 u thick, composed of a single layer. Micropyle absent. Sporulated oocysts with a polar granule. Oocyst residuum consists of a large globule resembling an oil droplet. Sporocysts ellipsoidal, without Stieda body, with a good deal of residual material between the sporozoites. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. GENUS Eimeria SCHNEIDER, 1875 4] Type Host: Spermophilus (syn., Citellus) lateralis (mantled ground squirrel). Location: Intestinal contents. Geographic Distribution: United States (Grand Canyon National Park, Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Only three oocysts were found in the intestinal contents of the host ground squirrel. The oocysts differed from those of all other coccidia described from Spermophilus and other rodents of the tribe Marmotini. They resembled those of Eimeria beckeri, but differed from them in having an oocyst polar granule and a single-layered rather than a double-layered wall. However, in view of the paucity of material available, the designation of this form as a new species does not seem warranted. HIMERITA™ (SPs EV TINE, 1952 Eimeria (?) sp. Levine, 1952(1951): 205-208. Description: Unsporulated oocysts 19-21 x 21-25 u, with a mean of 20.4 x 23.3 u; length-width ratio 1.1-1.2, with a mean of 1.1. Oocyst wall smooth, composed of two layers, the inner one dark yellowish and the outer colorless. Micropyle absent. No sporulated oocysts were seen. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) parryi (Parry’s ground squirrel). Location: Feces. Geographic Distribution: Canada (Arctic coast of Canadian Northwest Territories). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This form was found in five of seven S. parryi. Remarks: Since no sporulated oocysts were seen, it is impossible to assign this form a specific name or even to be positive of its genus. How- ever, it closely resembles Eimeria beckeri and may even belong to this species. EIMERIA VILASI DORNEY, 1962 (Plate 6, Fig. 55) Eimeria vilasi Dorney, 1962: 258-261. Description: Oocysts subspherical, occasionally ellipsoidal or ovoid. Two hundred sporulated oocysts measured 11-23 x 7-19 u, with a mean 42 THE COCCIDIAN PARASITES OF RODENTS of 18x14 u; their mean length-width ratio was 1.23. Oocyst wall com- posed of two layers: the outer one smooth, yellow-green, 0.7 u thick; the inner one dark tan, 0.3 uw thick. (Dorney said that the total thickness was 0.7-1.3 u.) Micropyle absent. Zero to six (mean, 1.5) oocyst polar granules present. Oocyst residuum absent. Sporocysts ellipsoidal (illus- trated as elongate ovoid); 25 sporocysts averaged 106 u. Stieda body small. Sporocyst residuum present. Sporozoites side by side in sporocysts, with a clear globule at larger end. Sporulation Time: Three to seven days at room temperature in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Tamias striatus (eastern chipmunk). Location: Primarily posterior quarter of small intestine (ileum). Geographic Distribution: North America (Wisconsin). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Dorney (1962) found this species in 99 per cent of 84 T. striatus in Vilas County, northeastern Wisconsin, and in south central Wisconsin. EIMERIA WISCONSINENSIS DORNEY, 1962 (Plate 6, Fig. 56) Eimeria wisconsinensis Dorney, 1962: 258-261. Description: Oocysts ellipsoidal. Fifty sporulated oocysts measured 26-35 x 20-27 u, with a mean of 31x24 u; their mean length-width ratio was 1.27. Oocyst wall composed of two layers: the outer one rough, yellow-tan, and 1.4-1.5 u thick; and the inner one colorless to pale pink and 0.3-0.4 u thick. Micropyle absent. One or rarely two oocyst polar granules present. Oocyst residuum absent. Sporocysts lemon-shaped; eight sporocysts measured 14—16 x 9-10 u, with a mean of 15x9 u. Stieda body prominent. Sporocyst residuum present, spread among and over sporo- zoites. Sporulation Time: Two to four weeks at room temperature in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Tamias striatus (eastern chipmunk). Location: Middle part of small intestine. Geographic Distribution: North America (Wisconsin). Pathogenicity: Unknown. Cross-Transmission Studies: None. GENUS Eimeria SCHNEIDER, 1875 43 Prevalence: Dorney (1962) found this species in 14 per cent of 84 T. striatus in Wisconsin. EIMERIA EUTAMIAE LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 7, Fig. 59) Eimeria eutamiae Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ovoid. Sixty sporulated oocysts measured 24-30 x 19-23 u, with a mean of 27.3x 21.4 u. Their length-width ratios ranged from 1.1—1.4, with a mean of 1.27. Oocyst wall smooth, composed of a very pale tan inner layer 0.4 uw thick and a colorless outer layer 1.0 u thick. The inner layer disappears at the small end of the oocyst. Micropyle absent. Sporulated oocysts with a polar granule which usually lies among the sporocysts. Oocyst residuum absent. Sporocysts lemon- shaped, about 11 x8 u, with a fairly thick transparent wall. Stieda body and sporocyst residuum present. Most sporocysts seem to le with their long axes perpendicular to the long axis of the oocyst, so that they are usually seen end on. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Eutamias dorsalis (cliff chipmunk). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one of two E. dorsalis in the Grand Canyon National Park, Arizona. EIMERIA ASIATICI N. SP. Eimeria beecheyi Henry. Vanabe and Okinami, 1940: 126-134. Description: Oocysts described as ovoid, but illustrated in the photo- micrographs as ellipsoidal, 15-22 10-12 u, according to Tanabe and Okinami (but the two oocysts illustrated in their photomicrographs were broader, measuring 17-19 14-15 u, if their stated magnification is correct). Oocyst wall colorless, composed of two layers, the outer one thick and the inner one thin. Micropyle absent. Oocyst residuum absent. Sporocyst residuum present, spherical. Sporulation Time: Three to four days. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Eutamias asiaticus (Asiatic chipmunk). Location: Cecal contents. Geographical Distribution: Japan (Keizyo). 44 THE COCCIDIAN PARASITES OF RODENTS Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described from two out of 24 E. asiaticus. Remarks: This species was reported by Tanabe and Okinami (1940) under the name Eimeria beecheyi, but it differs from that species in having a sporocyst residuum and also in occurring in Eutamias instead of Spermophilus. The only other species of Eimeria reported from Eutamias is E. eutamiae Levine, Ivens, and Kruidenier, 1957, from Eutamias dorsalis in the United States. The form from E. asiaticus is much smaller than this and is more ellipsoidal. ‘Tanabe and Okinami’s (1940) description is not complete enough to know whether there are other differentiating characters, but the oocysts in their photomicrographs do not resemble those of E. eutamiae. Fifteen other species of Ezmeria have been described from rodents of the tribe Marmotini. Five are from Marmota, one from Cynomys, and nine from Spermophilus. The form from Eutamias astaticus differs from E. bilamellata, E. volgensis, E. cynomysis, E. marmotae, E. arctomysi, and E. os, among other characters, in lacking a micropyle. It differs from E. lateralis, E. callospermophili, E. citelli, E. franklini, E. monacis, and E. perforoides, among other characters, in lacking an oocyst residuum. It differs from E. hoffmeisteri in having a two-layered wall and more elongate oocysts. It resembles E. beckeri more than any other species, but differs from it in having a colorless wall and more elongate oocysts. In view of these differences in morphology and host genus, it is con- sidered best to assign the form from E. asiaticus to a new species, Eimeria asiaticl nN. sp. Host Suborder SCIUROMORPHA Host Superfamily SCIUROIDEA Host Family SCIURIDAE Host Subfamily PETAURISTINAE EIMERIA GLAUCOMYDIS ROUDABUSH, 1937 * (Plate 4, Fig. 45) Eimeria glaucomydis Roudabush, 1937: 107-108. Description: Oocysts ellipsoidal, 18.5-12.3 x 13.2-10.6 u, with a mean of 16.2 11.5 u. Oocyst wall smooth, number of layers not stated. Micro- pyle absent. Oocyst residuum absent. Oocyst polar granule not figured, presumably absent. Sporocysts ellipsoidal, without Stieda body. Sporocyst residuum present. * The genitive of mys is myis, not mydis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 45 Sporulation Time: Slightly less than 20 hours. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Glaucomys volans (=volens) (flying squirrel). Location: Intestine. Geographic Distribution: United States (Iowa). Pathogenicity: Unknown. EIMERIA PARASCIURORUM BOND AND BOVEE, 1957 (Plate 6, Fig. 57; Plate 7, Figs. 61-65) Eimeria parasciurorum Bond and Bovee, 1957: 225-229. Eimeria sciurorum Galli-Valerio. Roudabush, 1937a: 107-108. [non] Eimeria sciurorum Galli-Valerio, 1922: 344-347. Description: Oocysts cylindrical, usually rounded at the ends but sometimes truncate. Fifty sporulated oocysts measured by Bond and Bovee (1957) in Florida were 22-36 x 12-20 u, with a mean of 29.0 x 16.0 u and a mean length-width ratio of 1.82. The oocysts measured by Rouda- bush (1937) in Iowa were 18-29 x 12-18 u, with a mean of 23.8 13.7 u. Oocyst wall smooth, light yellow-brown (according to Bond and Bovee) and 0.4—-0.6 u thick. Bond and Bovee stated that there is a dual mem- brane but illustrated the wall as composed of a single layer. Roudabush illustrated it similarly, but did not state the number of layers present. Micropyle absent. Oocyst polar granule and oocyst residuum absent. Sporocysts described by Bond and Bovee as egg-shaped and rounded at the end, but illustrated both by these authors and by Roudabush as ellipsoidal. Sporocysts 8-13 x 4-7 u, with a mean of We? 6: Ze ame ea mean length-width ratio of 1.81. Stieda body absent. Sporocyst residuum present. Sporozoites described by Bond and Bovee as piriform, but il- lustrated both by these authors and by Roudabush as elongate. Sporo- zoites 7-11 x 2-4 u, with a mean of 10.0 3.2 u and a mean length-width ratio of 3.11, according to Bond and Bovee. Sporulation Time: Twenty-two to thirty-six hours according to Bond and Bovee, twenty-eight hours according to Roudabush. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Glaucomys volans (=volens) (flying squirrel). Location: Intestine. Geographic Distribution: United States (Iowa, Florida). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in a female flying squirrel and a litter of several young at Gainesville, Florida, and in a single flying squirrel captured at Ames, Iowa. 46 THE COCCIDIAN PARASITES OF RODENTS Remarks: Roudabush (1937a) first described this species, but assigned it to Eimeria sciurorum Galli-Valerio, 1922, which had been described from the common squirrel, Scrurus vulgaris. In redescribing it, Bond and Bovee (1957) pointed out a number of morphological differences from E. scturorum and assigned it the new name, EF. parasciurorum. The fact that the hosts of these two species belong to different subfamilies gives additional justification, if such were necessary, for separating them. EIMERIA DORNEYI N. SP. Eimeria sp. Dorney, 1962: 258-261. Description: Oocysts ellipsoidal, a few truncate at one end, colorless to pale pink; 100 sporulated oocysts measured 17-30 x 10-19 u, with a mean of 2615 u; their mean length-width ratio was 1.7. Oocyst wall composed of a single layer, presumably smooth, light green under oil immersion. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts piriform; six sporocysts averaged 146 u. Stieda body present, not nipple-shaped. Sporocyst residuum present, usually dispersed among sporozoites. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Glaucomys sabrinus macrotis (northern flying squirrel). Location: Cecal contents. Geographic Distribution: North America (Wisconsin). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Dorney (1962) found this species in a single G. s. macrotis. Remarks: Dorney (1962) pointed out that this species differs morpho- logically from other species of Ezmeria described from Glaucomys, and that it fitted the description of EF. sciturorum from Sciurus niger. However, the description of E. sciwrorum is incomplete; moreover, Sczurus belongs to a different subfamily than Glaucomys. In view of the failure of all cross-transmission experiments of rodent Eimeria from one genus to another, it appears justifiable to assign the form from G. sabrinus a new name. We are therefore calling it Eimeria dorneyi n. sp. EIMERIA PETAURISTAE RAY AND SINGH, 1950 (Plate 7, Figs. 66-69) Eimeria petauristae Ray and Singh, 1950: 65-70. Description: Unsporulated oocysts flask-shaped, with a short neck and a dome-shaped “pseudomicropyle,” which forms a transparent cap at the anterior end and which disappears, becoming concave, on sporulation. GENUS Eimeria SCHNEIDER, 1875 47 Oocyst wall composed of two layers; the outer one rugged, deep brown, and 3.75—-6.25 u thick; the inner one thin, smooth, and transparent. The outer layer is readily broken away from the inner one by pressure. Oocysts measure 46.5—52.5 x 35.0-40.0 u. Sporocysts naviculoid, with one end slightly broader than the other, but without Stieda body. Sporocysts 25.5-31.25 x 8.75-10.0 u. Each sporocyst contains two club-shaped sporo- zoites with a vacuole at either end. Small oocyst residuum present. Oocyst polar granule absent. Sporocyst residuum present. The oocysts are heavier than those of most coccidia, since they cannot be floated up by centrifugation in sugar, salt, or zinc sulfate solutions. Sporulation Time: Ten to twelve days in potassium dichromate solu- tion. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Petaurista petaurista (syn., P. inornatus) (Himalayan flying squirrel). Location: Alimentary canal (found in feces). Geographic Distribution: India (Mukteswar). Pathogenicity: Unknown. Cross-Transmission Studies: Ray and Singh (1950) failed to infect local Indian rabbits with this species. Host Suborder SCIUROMORPHA Host Superfamily GEOMYOIDEA Host Family GEOMYIDAE Host Subfamily GEOMYINAE EIMERIA GEOMYDIS SKIDMORE, 1929 * (Plate 7, Fig. 60) Eimeria geomydis Skidmore, 1929: 183-184. Description: Oocysts spherical to slightly ovoid, 12-15 x 12-13 u, with a mean of 13.3 x 12.5 wand a mean length-width ratio of 1.07. Oocyst wall smooth, colorless, 0.5 u thick, composed of two layers. (Only a single layer appears in Skidmore’s drawings.) Micropyle seen in only a few oocysts. Sporont almost fills unsporulated oocysts. Oocyst polar granule and oocyst residuum absent. Sporocysts 5—7 x 4-5 u, without Stieda body but with sporocyst residuum. Sporulation Time: Four days at room temperature in 2% potassium dichromate. * The genitive of mys is myis, not mydis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 48 THE COCCIDIAN PARASITES OF RODENTS Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Geomys bursarius (pocket gopher). Location: Small and large intestines. Geographic Distribution: United States (Nebraska). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was described from a single pocket gopher in Lincoln, Nebraska. EIMERIA THOMOMYSIS LEVINE, IVENS, AND KRUIDENIER, Iie (Plate 8, Fig. 70) Eimeria thomomysis Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts spherical to subspherical. Oocyst wall composed of a single layer, smooth, pale yellowish brown, 0.8 u thick, lined by a thin membrane. Micropyle absent. Nineteen sporulated oocysts from two host animals measured 13-16 x 13-16 u, with a mean of 14.2 x 13.9 u. Their length-width ratios ranged from 1.0-1.1, with a mean of 1.01. Oocyst polar granule and oocyst residuum absent. Sporocysts 106 u, with a small Stieda body and a few scattered residual granules. Sporo- cysts almost fill oocyst. Sporozoites lie head to tail in sporocyst and con- tain a large, spherical, clear, colorless body about 2 u in diameter at the large end. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Thomomys bottae (pocket gopher). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in two out of five T. bottae from Grand Canyon National Park, Arizona. Host Suborder SCIUROMORPHA Host Superfamily GEOMYOIDEA Host Family HETEROMYIDAE Host Subfamily PEROGNATHINAE * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 49 EIMERIA PEROGNATHI LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 8, Fig. 71) Eimeria perognathi Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ovoid. Oocyst wall composed of a single layer somewhat rough, yellowish brown, 1 u thick. Micropyle absent. Five sporulated oocysts measured 19-22 x 15-16 u, with a mean of 20.2 x 15.3 u. Their length-width ratios ranged from 1.2-1.4, with a mean of 1.32. Oocyst polar granule absent. Oocyst residuum composed mostly of large, clear globules. Sporocysts 6-7 x 4-5 u, with a small, flat Stieda body and a small amount of scattered residual material. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Perognathus intermedius (rock pocket mouse). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of two P. intermedius from the Grand Canyon National Park, Arizona. EIMERIA PENICILLATI IVENS, KRUIDENIER, AND LEVINE, 1959 (Plate 8, Fig. 78) Eimeria penicillati Ivens, Kruidenier, and Levine, 1959: 53-57. Description: Oocysts subspherical, ellipsoidal, or slightly ovoid. Oocyst wall pale brownish yellow or tan, smooth, composed of a single layer (confirmed by breaking the oocyst) about 0.6 u thick. Micropyle absent. Six sporulated oocysts from the type host (Perognathus penicillatus) measured 16-20 x 14-16 u, with a mean of 17.8x 14.7 u. Their length- width ratios ranged from 1.1—-1.3, with a mean of 1.18. Sporocysts broadly lemon-shaped, 9x7 u, with a length-width ratio of 1.33. Stieda body small, rounded. Oocyst residuum composed of one to several large, clear globules. Oocyst polar granule present. Sporocyst residuum com- posed of a number of large granules. Sporozoites oriented more or less lengthwise in sporocysts. Three oocysts from a P. flavus measured 17-20 x 15-19 u. They differed from those of P. penicillatus only in having either one or two polar granules instead of a single one. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: Not described. 50 THE COCCIDIAN PARASITES OF RODENTS Type Host: Perognathus penicillatus (pocket mouse). Other Host: Perognathus flavus (pocket mouse). Location: Intestinal contents. Geographic Distribution: Mexico (Sonora, Chihuahua). Pathogenicity: Unknown. Cross-Transmission Studies: None. Host Suborder SCIUROMORPHA Host Superfamily GEOMYOIDEA Host Family HETEROMYIDAE Host Subfamily DIPODOMYINAE EIMERIA MOHAVENSIS DORAN AND JAHN, 1959 (Plate 8, Figs. 72-77; Plate 9, Figs. 79 and 80) Eimeria mohavensis Doran and Jahn, 1949: 631-632; Doran and Jahn, 1952: 93-101; Doran, 1953: 31-60. Description: Oocysts ellipsoidal. Oocyst wall composed of a single layer, smooth, light brown, 0.7—0.9 u thick. Micropyle absent. Sporulated oocysts 22-26 x 14-18 u, with a mean of 24.1 15.7 wu and a mean length-width ratio of 1.54. Sporont in unsporulated oocysts 12-15 x 12-16 u, with a mean of 13.6 x 12.7 u. Oocyst polar granule and oocyst residuum absent. Doran and Jahn (1952) gave the sporocyst dimensions as 6-10 x 5.5-8 u, with a mean of 7.87.6 u, but in their illustration (their Fig. 10) the sporocysts appear to be about 7 x 4 u. Sporocysts without Stieda body, but with a residuum 4.0—4.5 u long. Sporulation Time: ‘Twenty-four to sixty hours in 2-4% potassium dichromate at the optimum temperature, 23-27C; 64-117 hours at 9.5C; 44—78 hours at 40C. No sporulation at 2.5 or 45C. Schizogony: Doran (1953) described the schizogonic part of the life cycle in detail. There are two sizes of schizonts and of merozoites. The small, early schizonts, 4.1-10.5 u in diameter, are present on the second through the fifth days of the prepatent period. ‘These produce 20 to 35 comma-shaped merozoites measuring 6.5-8.5 x 1.5-2.0 u, with a central nucleus. The larger, later schizonts, 8.0-10.0 uw in diameter, are present on the sixth and seventh prepatent and first and second patent days. They produce 50 to 75 merozoites measuring 4.5—-6.5 x 1.0-1.5 u, with a central nucleus. Since the first gametes and gametocytes appear on the fifth prepatent day, one day before the first large schizonts, it is clear that some of the first-generation merozoites must produce gametes and gametocytes directly. Most, however, probably produce second-generation schizonts. Assuming that 33 of the 35 merozoites of the small schizonts developed into large schizonts, that 1 of these merozoites and | of those GENUS Eimeria SCHNEIDER, 1875 51 produced by each large schizont developed into a microgametocyte, and that 1 of the small schizont merozoites and 74 of the large schizont merozoites developed into macrogametes, Doran (1953) calculated that the theoretical maximum yield of oocysts from an initial infection of 100 oocysts should be 1,954,400. He actually obtained an average oocyst yield of 9,710,000. It is obvious, therefore, that there must be more than two generations of schizonts and merozoites. Since the small schizonts are present for a longer time than the large ones, Doran considered that the additional generation or two is probably of this type. On the basis of his work, he considered the most likely pattern to be that some of the merozoites produced by the first-generation, small schizonts develop di- rectly into gametes and gametocytes, while others develop into second- generation small schizonts and still others into second-generation large schizonts. The merozoites produced by the second-generation small schizonts then develop into third-generation large schizonts and probably also into gametes. The merozoites from both the second- and_third- generation large schizonts probably develop only into gametocytes and gametes. Further study will be needed, however, to verify this scheme. Gametogony: This has been described by both Doran and Jahn (1952) and Doran (1953). Gametocytes and gametes are found during the last two days of the prepatent and first to sixth days of the patent periods. ‘The macrogametes are round and measure 10-16 uw in diameter. The microgametocytes are ellipsoidal, 17.0-18.5 x 21.0-22.5 u, and produce a large number of microgametes. Prepatent Period: The prepatent period following experimental infec- tion is seven days. The patent period, during which oocysts are present in the feces, lasts nine to ten days. Type Host: Dipodomys panamintinus mohavensis (kangaroo rat). Other Hosts: No other natural hosts have been reported, but Doran (1953) produced experimental infections in Dipodomys merriami, D. nitratoides, D. heermanni, D. deserti, and D. agilis. Location: The small schizonts are found throughout the small in- testine and cecum, most being present in the distal part of the small intestine. The large schizonts are more prevalent in the cecum than in the small intestine. The gametes and gametocytes are found in the cecum. All stages are above the nuclei of the host epithelial cells. Geographic Distribution: United States (southern California). Pathogenicity: Unknown. Cross-Transmission Studies: Doran (1951, 1953) successfully infected three other subspecies of Dipodomys panamintinus, D. p. leucogenys, D. p. panamintinus, and D. p. caudatus. He also infected 11 of 12 D. merriami merriamt, 11 of 12 D. nitratoides brevinasus, 5 of 6 D. heermanni mor- or nN THE COCCIDIAN PARASITES OF RODENTS roensis, all of 6 D. h. tularensis, 1 D. h. swarthi, 1 D. deserti deserti, and 6 D. agilis agilis. He was unable to infect 8 Perognathus longimembris, 7 P. formosus mohavensis, 4 Peromyscus boylit, 12 P. maniculatus, 12 P. cali- fornicus, 7 P. truei, 5 Onychomys torridus, 4 Neotoma lepida, 8 Spermo- philus leucurus, 4 Mus musculus, and 4 Rattus norvegicus. Thus this species appears confined to the genus Dipodomys. It is of interest in this respect that, although Dipodomys merriami merriami was readily infected in the laboratory, natural infections were not found in any of 197 D. m. merriami trapped in the field. This was despite the fact that D. m. merriami is sympatric with D. p. mohavensis, that they have similar food habits and forage over the same range, that they have been taken in the same trap within half an hour, and that D. m. merriami frequently enters the burrows of other kangaroo rats, including presumably D. p. mohavensis. Prevalence: E. mohavensis was found by Doran (1953) in 8.7 per cent of 251 Dipodomys panamintus mohavensis in southern California. EIMERIA DIPODOMYSIS LEVINE, IVENS, AND KRUIDENIER, 1958 * (Plate 10, Fig. 81) Eimeria dipodomysis Levine, Ivens, and Kruidenier, 1958: 291-298. Description: Oocysts ellipsoidal. Oocyst wall yellowish brown, rough, composed of two layers, the outer one 3.5 u thick at the sides and 3.0 u thick at the ends, the inner one 0.7 u thick. Micropyle absent. Seventeen sporulated oocysts measured 47-61 x 38—42 u, with a mean of 54.1 x 40.2 u. Their length-width ratios ranged from 1.2-1.5, with a mean of 1.35. Sporocysts ovoid, about 16x11 u. Stieda body shaped somewhat like a conical dome, i.e., excavated in the center. No oocyst polar granule could be distinguished. Oocyst residuum large, composed of a mass of homoge- neous granules. In some oocysts these granules are few, large, and loosely aggregated, while in others they are small and numerous, forming a compact mass. Sporozoites embedded in a large amount of sporocyst residual material. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dipodomys phillipst (Mexican kangaroo rat). Location: Intestinal contents. Geographic Distribution: Mexico (Aguascalientes). Pathogenicity: Unknown. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 53 Cross-Transmission Studies: None. Prevalence: This species was found in one out of three D. phillipsi from Aguascalientes and Jalisco, Mexico. Host Suborder SCIUROMORPHA Host Superfamily GEOMYOIDEA Host Family HETEROMYIDAE Host Subfamily HETEROMYINAE EIMERIA LIOMYSIS LEVINE, IVENS, AND KRUIDENIER, 1958 * (Plate 10, Fig. 83) Eimeria liomysis Levine, Ivens, and Kruidenier, 1958: 291-298; Ivens, Krui- denier, and Levine, 1959: 53-57. Description: Oocysts subspherical to ellipsoidal. Oocyst wall composed of two layers: the outer one 0.9 u thick, pale yellow, slightly rough and pitted; and the inner one 0.3 u thick and practically colorless. Micropyle absent. Seventy-five sporulated oocysts from two specimens of the type host (Liomys pictus) measured 15-24 x 14-21 u, with a mean of 19.5 x 17.7 u. Their length-width ratios ranged from 1.0-1.3, with a mean of 1.10. Forty-four sporulated oocysts from a Liomys irroratus measured 15-23 x 14-20 u, with a mean of 18.2 17.2 u. Their length-width ratios ranged from 1.0-1.1, with a mean of 1.06. Eight sporulated oocysts measured by Ivens, Kruidenier, and Levine (1959) from two L. pictus measured 18-23 x 17-21 u, with a mean of 20.8x 19.1 u and a mean length-width ratio of 1.07. Sporocysts almost ellipsoidal to ovoid, about 107 u, with a small Stieda body. Oocyst polar granule present. Oocyst residuum absent. Sporozoites usually at the ends of the sporocysts, with some relatively large residual granules between them. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Liomys pictus (painted spiny pocket mouse). Other Host: Liomys irroratus (Mexican spiny pocket mouse). Location: Intestinal contents. Geographic Distribution: Mexico (Sinaloa, Jalisco, Nayarit). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found by Levine, Ivens, and Kruidenier (1958) in two L. pictus from Sinaloa, Mexico, and in one L. trroratus from Jalisco, Mexico. It was found by Ivens, Kruidenier, and Levine (1959) in a L. pictus from Nayarit, Mexico, and another from Sinaloa. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of the name, so the error must be perpetuated. 54 THE COCCIDIAN PARASITES OF RODENTS EIMERIA PICTI LEVINE, IVENS, AND KRUIDENIER, 1958 (Plate 10, Fig. 84) Eimeria picti Levine, Ivens, and Kruidenier, 1958: 291-298. Description: Oocysts subspherical to ellipsoidal. Oocyst wall composed of two layers: the outer one brownish yellow, 1.3 u thick, rough, and pitted; and the inner one brownish yellow and 0.4 u thick. When the outer wall is broken, the inner wall usually remains intact. Micropyle absent. Fifty-three sporulated oocysts measured 22-32 x 19-28 u, with a mean of 25.8 x 22.5 u. Their length-width ratios ranged from 1.0-—1.2, with a mean of 1.15. Sporocysts broadly lemon-shaped, 11-12 x 8-9 u. Stieda body present. One to two oocyst polar granules present. Oocyst residuum usually composed of a number of large, clear, irregular gran- ules, but sometimes a single, large, granular mass. Sporocyst residual material composed of a few to many granules. Sporozoites lie longi- tudinally, head to tail, in the sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Liomys pictus (painted spiny pocket mouse). Location: Intestinal contents. Geographic Distribution: Mexico (Sinaloa). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one of two L. pictus from Sinaloa, Mexico. Host Suborder SCLUROMORPHA Host Superfamily CASTOROIDEA Host Family CASTORIDAE Host Subfamily CASTORINAE EIMERIA SPREHNI YAKIMOFF, 1934 Eimeria sprehni Yakimoff, 1934: 294. Description: Oocysts ovoid (ellipsoidal in drawing), sometimes with one side concave. Oocyst wall “doppelt konturiert,” but illustrated as composed of a single, apparently smooth layer. Micropyle absent. ‘Thirty- five oocysts measured 16-20 x 11-14 u, with a mean of 17.6 x 12.0 u. Their length-width ratios ranged from 1.3-1.7, with a mean of 1.47. Oocyst polar granule and oocyst residuum present. Sporocysts elongate, ap- parently without Stieda body. Complete sporulation did not take place, and no sporozoites were seen. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. i 3 ions GENUS Eimeria SCHNEIDER, 1875 Type Host: Castor canadensis (Canadian beaver). Location: Feces. Geographic Distribution: USSR (presumably in a zoo). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff (1934) found this species in one of three Canadian beavers in Voronezh. Remarks: Although both Becker (1956) and Pellérdy (1956) listed Castor fiber as a host of this species in addition to C. canadensis, we have not found a report of it in the former host. Yakimoff (1934) ex- amined 19 C. fiber from Russia without finding coccidia. His discussion, however, was misleading. He first stated that he examined “22 Bibern (Castor fiber), and then went on to say that he found no coccidia in 19 from Voronezh, but did find a few coccidia in 1 out of 3 ‘‘kanadischer Biber.” This must have been C. canadensis, although he did not use this scientific name in his paper. Yakimoff (1934) also stated that Sprehn (1932) found coccidia in “bibern (Sumpfbiberraten),” but the Sumpfbi- berrat is the nutria (Myocastor coypus). Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily CRICETINAE Host Tribe HESPEROMYINI EIMERIA ORYZOMYSI CARINI, 1937 * (Plate 36, Figs. 295-297) Eimeria oryzomysi Carini, 1937a: 47-49; Carini, 1937b: 624-627. Description: Oocysts ellipsoidal to ovoid. Oocyst wall with a double contour (illustrated as a single layer), smooth, light brown. Oocysts 22-25 x 17-19 u, with a length-width ratio of 1.25. Micropyle absent. In freshly passed oocysts, the sporont completely fills the oocyst, but it shrinks away from the wall as it develops. Oocyst polar granule apparently absent. Oocyst residuum at first 8-10 u in diameter and granular, but as the sporocysts develop the granules tend to coalesce so that frequently a single clear globule of varying size results. Sporocysts ovoid, 11 x8 u, with a Stieda body and a sporocyst residuum composed of centrally located small, scattered gran- ules. Sporozoites lie lengthwise in sporocysts. * The genitive of mys is myis, not mysi. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 56 THE COCCIDIAN PARASITES OF RODENTS Sporulation Time: Five to six days at 20-25C. Schizogony: Schizogony takes place in the epithelial cells of the small intestine and especially of the duodenum. As the schizonts develop, they push the host cell nucleus aside and may come to lie below it. The schizonts form 15 to 20 banana-shaped merozoites measuring about 8 x 2-2.5 u. The merozoites encountered free in the lumen of the in- testine measured 10-11] x 3 u. The above description is based on stages found 6 and 12 days after experimental infection. Gametogony: Gametogony takes place in the small intestine. Carini (1937a,b) illustrated macrogametes and microgametocytes both above and below the host cell nuclei. The macrogametes have a central nucleus and a border of hematoxylin-staining plastic granules which disappear when the oocyst membrane is formed. The microgametocytes form a large number of microgametes 2-2.5 u long. Prepatent Period: Five days. Type Host: Oryzomys sp. (rice rat). Location: Small intestine. Geographic Distribution: Brazil (Sao Paulo). Pathogenicity: Unknown. One of two experimentally infected rice rats died on the twelfth day without previous signs of illness. Cross-Transmission Studies: Carini (1937a,b) was unable to infect white or common mice with this species. EIMERIA COUESIT KRUIDENIER, LEVINE, AND IVENS, 1960 (Plate 10, Fig. 82) Eimeria couesti Kruidenier, Levine, and Ivens, 1960: 100-101. Description: Oocysts ellipsoidal, pale yellowish. Oocyst wall somewhat rough and pitted, heavy, composed of a single layer about 1.3 u thick with weak radial striations. Five sporulated oocysts measured 20-23 x 17-20 u, with a mean of 21.4x 18.0 u; their length—width ratio was 1.20. Micropyle absent. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid. Seven sporocysts measured 10-14 x 7-8 u, with a mean of 11.7 7.7 u; their length-width ratios ranged from 1.2—-1.8, with a mean of 1.53. Stieda body present. Sporozoites oriented longitudinally in sporocysts, with what appears to be a large, clear, yellowish globule at the large end. Sporocyst residuum composed of some rather loose granules or absent. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: Not described. Type Host: Oryzomys c. couesi (rice rat). Location: Feces. GENUS Eimeria SCHNEIER, 1875 57 Geographic Distribution: Mexico (Oaxaca). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA PEROMYSCI LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 11, Fig. 85) Eimeria peromysci Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ellipsoidal. Oocyst wall composed of two layers; the outer one 1.3 u thick, yellowish brown, and rough; the inner one 0.4 u thick and yellowish brown. Micropyle absent. Nineteen sporulated oocysts measured 26-32 x 21-27 u, with a mean of 29.4 23.9 u. Their length-width ratios ranged from 1.1-1.3, with a mean of 1.22. Sporulated oocysts with an oocyst polar granule and one to several large, clear, colorless residual globules up to 10 uw in diameter. Sporocysts about 15x9 u, lemon-shaped, with a Stieda body at one end. Sporocyst wall about 0.4 u thick. Sporocysts contain two sporozoites lying with the large end of one beside the small end of the other. A large number of coarse residual granules fill the space between the sporozoites. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus truei (pinon mouse). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of four P. true: in Grand Canyon National Park, Arizona. EIMERIA ARIZONENSIS LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 11, Figs. 87 and 88; Plate 12, Fig. 89) Eimeria arizonensis Levine, Ivens, and Kruidenier, 1957: 80-88; Levine and Ivens, 1960: 207-212. Description: This species has been reported from three species of Peromyscus. Since there are minor differences in the appearance of the oocysts from the different hosts, each form is described separately. Form from type host, Peromyscus truei: Oocysts subspherical to el- lipsoidal. Oocyst wall smooth, pale tan, composed of a single layer about 0.9 u thick, lined by a thin membrane. Micropyle absent. Fifty sporulated oocysts from one host animal from the north rim of Grand Canyon measured 19-27 x 17-22 uw, with a mean of 21.7x19.2 u. Their length- width ratios ranged from 1.0-1.2, with a mean of 1.13. Eight sporulated 58 THE COCCIDIAN PARASITES OF RODENTS oocysts from another host animal from the south rim of Grand Canyon measured 22-29 x 18-23 u, with a mean of 25.2x 20.7 u. Their length- width ratios ranged from 1.1—1.3, with a mean of 1.22. Sporulated oocysts with an oocyst polar granule and usually with a single large, clear, colorless residual globule about 4 u in diameter. Sporocysts almost fill oocyst. Sporocysts about 117 u, lemon-shaped, with a Stieda body at one end. Sporocysts contain two sporozoites lying with the large end of one beside the small end of the other. Sporocyst residuum more or less compact, composed of coarse granules. Form from P. maniculatus: Oocysts ellipsoidal. Oocyst wall light yel- lowish, slightly to moderately pitted, occasionally smooth, composed of a single layer (confirmed by crushing the oocyst) 1.3-1.7 u thick. Micro- pyle absent. Forty sporulated oocysts measured 22-29 x 18-23 u, with a mean of 26.1 x 20.8 u; their length-width ratios ranged from 1.1—1.3, with a mean of 1.24. Oocyst polar granule present. Oocyst residuum composed of a cluster of large, homogeneous granules or globules 2—5 u in diameter. Sporocysts lemon-shaped, with prominent Stieda body. Sporocyst wall 0.4 u thick. Twenty-three sporocysts measured 11-13 x 7-9 u with a mean of 12.2x8.0 u; their length-width ratios ranged from 1.3-1.7, with a mean of 1.56. Sporozoites homogeneous, colorless, lying lengthwise in sporocysts. Sporocyst residuum composed of coarse granules which fill the space between the sporozoites. Form from P. lewcopus: Oocysts ellipsoidal to broadly ellipsoidal. Oocyst wall pale yellowish to yellowish, occasionally almost smooth but usually more or less pitted, composed of a single layer (confirmed by crushing the oocyst) 1.0-1.2 u thick. Micropyle absent. Sixty-five spor- ulated oocysts from two host individuals measured 20-25 x 17-21 u, with a mean of 22.4x 19.4 u; their length-width ratios ranged from 1.1—1.3, with a mean of 1.16. One or two oocyst polar granules present. Oocyst residuum present, usually a single, large, waxy-appearing globule about 4 w in diameter. Sporocysts lemon-shaped to rather ovoid, with prominent Stieda body. Sporocyst wall rather thick. ‘Twenty-two sporocysts meas- ured 12-14 x 7-8 u, with a mean of 12.6 7.2 u; their length-width ratios ranged from 1.6-1.9, with a mean of 1.75. Sporocyst residuum composed of more or less scattered coarse granules. Sporozoites colorless, lying lengthwise, head to tail, in sporocysts. Schizogony and Gametogony: Form from P. truet: Unknown. Form from P. maniculatus: A few microgametocytes, macrogametes, and young oocysts were seen by Levine and Ivens (1960) in the epithelial cells of the villi of the anterior ileum. The macrogametes had a single layer of eosinophilic plastic granules. Their relation to the host cell nucleus was uncertain. They measured approximately 19x14 uw when GENUS Eimeria SCHNEIDER, 1875 59 mature, but were shrunken. The young oocysts measured approximately 2216 u. A few macrogametes were also seen in the posterior ileum. No endogenous stages of this form were found in the duodenum, jejunum, or cecum. Form from P. leucopus: A moderate number of macrogametes, micro- gametocytes, and oocysts were found by Levine and Ivens (1960) in the epithelial cells of the tips and sides of the villi, and very rarely in the crypts, of the jejunum. Fewer parasites were found in the anterior ileum, still fewer in the posterior ileum, and only a single macrogamete was seen in the duodenum. No endogenous stages were seen in the cecum. All stages were present both above and below the host cell nuclei; the older forms tended to lie below them. The macrogametes ranged in size up to about 14x 10 u, and had a single layer of eosinophilic plastic granules. The microgametocytes ranged in size up to about 15x8 u. The oocysts ranged in size up to about 19x 14 u. A very few schizonts were seen in the jejunum, and a single one was found in a detached piece of villus in the posterior ileum. They lay beneath the host cell nuclei in the epithelial cells at the sides of the villi, but too few were seen to be sure that this is their only location. They measured 9-12 x 7-9 u. Most seemed to contain about 8 or perhaps a few more sausage-shaped merozoites about 1 u in diameter, but one schizont contained perhaps 16 to 24; these may possibly have belonged to two different generations. There was no schizont residuum. Prepatent Period: Unknown. Type Host: Peromyscus truei (pinon mouse). Other Hosts: Peromyscus maniculatus, P. leucopus (deer mice). Location: Form from P. truez: Intestinal contents. Form from P. maniculatus: Anterior and posterior ileum. Form from P. leucopus: Jejunum, anterior ileum, posterior ileum, rarely duodenum. Geographic Distribution: United States (Arizona, Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine, Ivens, and Kruidenier (1957) found this species in two of four P. truei in Grand Canyon National Park, Arizona. Levine and Ivens (1960) found it in one of nine P. maniculatus and two of seven P. leucopus from the vicinity of Sullivan, Ilinois. Remarks: Although there was a significant difference in size and shape of the oocysts from the two P. true, they were not sufficient to merit taxonomic separation. It is of interest that one of these pinon mice came from the north rim of Grand Canyon and the other from the south rim. Both were P. t. trwei, derived from the same general invading popula- tion. This subspecies lives in juniper rock habitats and does not occur in 60 THE COCCIDIAN PARASITES OF RODENTS the depths of Grand Canyon. However, according to D. F. Hoffmeister, intercommunication was highly probable during the Pleistocene and is possible even now. The form from P. maniculatus differs from that from P. true: only in that its oocyst wall is thicker and is ordinarily more or less pitted, in that it lacks a membrane lining the oocyst wall, and in that its oocyst residuum is composed of a cluster of homogeneous globules rather than a single large one. The form from P. leucopus differs from that from P. truei only in that its oocyst wall is more or less pitted and in that it lacks a membrane lining the oocyst wall. The oocysts of the form from P. leucopus differ from those of the form from P. maniculatus only in having a thinner oocyst wall and in that the oocyst residuum is com- posed of a single, large, homogeneous globule rather than a cluster of them. Those endogenous stages of the forms from P. maniculatus and P. leucopus which were seen did not appear to differ morphologically. However, none was seen in the jejunum of P. maniculatus, while they were most common in this location in P. leucopus. Since so few were found in the former, this should not be considered a valid difference. The above differences do not appear sufficient to justify the establishment of new species, although future research may reveal differences in the endogenous stages, location in the host, or host-parasite relations which would do so. EIMERIA EREMICI LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 12, Fig. 92) Eimeria eremici Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ellipsoidal. Oocyst wall smooth, composed of two layers, the outer one | u thick and colorless, and the inner one 0.4 u thick and pale tan. Micropyle absent. Fifty sporulated oocysts from one host animal measured 22-30 x 18-22 u, with a mean of 25.3x 20.8 u. Their length-width ratios ranged from 1.1-1.4, with a mean of 1.21. Sporulated oocysts with one to three polar granules and a single, clear residual globule about 6-8 u in diameter. (In 3 out of 50 oocysts these oocyst residua were granular rather than clear and homogeneous.) Sporocysts about 108 u, lemon-shaped, with a relatively inconspicuous Stieda body at one end. The two sporozoites le lengthwise in the sporocysts, which contain more or less coarsely granular residual material. Schizogony: Unknown. Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus eremicus (cactus mouse). Location: Intestinal contents. GENUS Eimeria SCHNEIDER, 1875 61 Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of two P. eremicus in Grand Canyon National Park, Arizona. EIMERIA LANGEBARTELI IVENS, KRUIDENIER, AND LEVINE, 1959 (Plate 11, Fig. 86) Eimeria langebarteli Ivens, Kruidenier, and Levine, 1959: 53-57. Description: Oocysts elongate ellipsoidal. Oocyst wall smooth, pale yellowish, composed of a single layer about 0.8 u thick. Micropyle absent. Fifteen sporulated oocysts from two host animals measured 20-23 x 13-14 u, with a mean of 21.0 13.6 u. Their length-width ratios ranged from 1.4-1.7, with a mean of 1.55. Sporocysts elongate ellipsoidal, thin-walled. Ten sporocysts measured 8-10 x 5-6 u, with a mean of 9.45.3 u; their length-width ratios ranged from 1.6—2.0, with a mean of 1.78. Stieda body small. Oocyst polar granule present. Oocyst residuum absent. Sporozoites elongate, curled inside sporocysts, with many small granules so closely adherent to the sporozoites that it was not possible to be sure whether they were inside or outside the sporozoite membrane. Hence the presence of true sporocyst residual material was not definitely established. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: Not described. Type Host: Peromyscus boylit (deer mouse). Location: Intestinal contents. Geographic Distribution: Mexico (Chihuahua). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Ivens, Kruidenier, and Levine (1959) found this species in two of four P. boylii from near Cuauhtémoc, Chihuahua, Mexico. EIMERIA CAROLINENSIS VON ZELLEN, 1959 (Plate 12, Fig. 91) Eimeria carolinensis von Zellen, 1959: 104-105. Description: Oocysts ellipsoidal, occasionally elongate ellipsoidal. Eighteen sporulated oocysts measured 14-19 x 10-13 u, with a mean of 17.6 x 11.3 u; their length-width ratios ranged from 1.2—-1.6, with a mean of 1.43. Micropyle absent. Oocyst wall smooth, composed of an outer colorless layer 1 u thick and an inner dark brown layer 0.5 wu thick. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, 62 THE COCCIDIAN PARASITES OF RODENTS almost filling oocyst. Sporocysts about 8.54.5 u, with a small Stieda body. Sporocyst residuum present on one side of the sporozoites, being partially concealed by them. Sporozoites lie lengthwise in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus leucopus (white-footed mouse). Location: Intestinal contents. Geographic Distribution: United States (North Carolina). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Von Zellen (1959) found this species in 17 per cent of 53 P. leucopus from the vicinity of Durham, North Carolina, between Janu- ary 12 and November 24, 1958. EIMERIA DELICATA LEVINE AND IVENS, 1960 (Plate 12, Fig. 90) Eimeria delicata Levine and Ivens, 1960: 207-212. Description: Oocysts ellipsoidal. Oocyst wall very pale yellowish, smooth, composed of a single layer about 0.6 u thick. Micropyle absent. Eleven sporulated oocysts measured 13-15 x 10-12 u, with a mean of 14.2 11.3 uw; their length-width ratios ranged from 1.1—1.4, with a mean of 1.25. Oocyst polar granule present. Oocyst residuum absent. Sporocysts elongate ovoid with a rather pointed end bearing a tiny Stieda body, very thin-walled, 8x4—-5 u, with a length-width ratio of about 1.9. Sporozoites oriented lengthwise in sporocysts. Some sporocyst residual granules ordinarily present, although they may sometimes be absent. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus maniculatus (deer mouse). Location: Intestinal contents. Geographic Distribution: United States (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine and Ivens (1960) found this species in two out of nine P. maniculatus from the vicinity of Sullivan, Illinois. EIMERIA ROUDABUSHI LEVINE AND IVENS, 1960 (Plate 12, Fig. 93) Eimeria roudabushi Levine and Ivens, 1960: 207-212. Description: Oocysts ellipsoidal. Oocyst wall smooth, almost colorless, composed of a single layer about 1.3 wu thick at the sides, grading to about 0.9 uw thick at the ends. Micropyle absent. “Twenty sporulated GENUS Eimeria SCHNEIDER, 1875 63 oocysts measured 20-26 x 17-20 u, with a mean of 22.2 18.6 u; their length-width ratios ranged from 1.1-1.3, with a mean of 1.19. Oocyst polar granule present. Oocyst residuum atypical, composed of a small amount of cobwebby-appearing material at both ends of the oocyst. Sporocysts ovoid, thin-walled, with a small Stieda body. Sporocysts 12-138 u with a length-width ratio of 1.6-1.7. Sporocyst residuum large, compact. Sporozoites colorless, lying lengthwise, head to tail, in sporocysts, but so long that they are folded back upon themselves. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus leucopus (deer mouse). Location: Intestinal contents. Geographic Distribution: United States (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine and Ivens (1960) found this species in one out of seven P. leucopus from the vicinity of Sullivan, Illinois. FIMERIA LEUCOPI VON ZELLEN, 1961 (Plate 13, Figs. 94-97; Plate 14, Figs. 98-101; Plate 15, Fig. 102) Eimeria leucopi von Zellen, 1961: 134-138. Description: Oocysts ellipsoidal, occasionally ovoid, rarely spherical. Oocyst wall uniformly thick, rough, composed of an outer, yellowish, transparent layer 0.5 uw thick and an inner, dark brown layer 1.0 u thick. Micropyle absent. Three hundred and fifty-nine oocysts measured 14-24 x 14-21 u, with a mean of 19x 17 u; their length-width ratios ranged from 1.0-1.6, with a mean of 1.1. Oocyst polar granule absent. Oocyst residuum usually composed of one to several clear, green globules up to 6 uw in diameter. Sporocysts 11-14 x 6-8 u, with a mean of 127 u. Stieda body present. Sporocyst residuum present, compact or diffuse. Sporozoites 4—7 x 2-3.5 u, with a mean of 5xX2.4 u lying lengthwise in sporocysts. Sporulation Time: Fifty-four to one hundred and twenty-eight hours on vaseline-ringed slides. Sporogony: Von Zellen (1961) described sporogony of these oocysts in S/4 sodium chloride solution on vaseline-ringed slides, presumably at room temperature. The sporont of newly discharged oocysts completely fills the oocysts. It contracts after 3 hours. About 26 hours later, clear areas, never entirely devoid of granules, appear within the sporont. About 50 hours after the oocysts have been discharged, a “fertilization spindle” appears. This is followed by the buckelbildung or ball stage, in which the limiting membrane of the sporont fades, the protoplasmic mass loses its dark yellowish brown color and becomes largely colorless, 64 THE COCCIDIAN PARASITES OF RODENTS and the clear area surrounding it changes in color from yellowish to faint blue. Small, homogeneous, green globules appear within the gran- ular portion of the sporont and quickly associate with the clear material of each developing bulge. This process of pyramid formation begins about 62 hours after oocyst discharge and lasts about 2.5 hours. Schizogony and Gametogony: Unknown. According to von Zellen (1961), the patent period is six days. Prepatent Period: Five to six days, according to von Zellen (1961). Type Host: Peromyscus leucopus (deer mouse). Location: Small intestine. Geographic Distribution: North America (North Carolina). Pathogenicity: Unknown. Cross-Transmission Studies: Von Zellen (1961) was unable to infect the golden deermouse, Peromyscus nuttalli, with E. leucopt. Prevalence: Von Zellen (1961) found E. leucopi in 13 per cent of 53 Peromyscus leucopus from the vicinity of Durham, North Carolina. EIMERIA BAIOMYSIS LEVINE, IVENS, AND KRUIDENIER, 1958 * (Plate 15, Fig. 103) Eimeria baiomysis Levine, Ivens, and Kruidenier, 1958: 291-298; Kruidenier, Levine, and Ivens, 1960: 100-101. Description: Oocysts ellipsoidal. Oocyst wall yellowish, quite rough and pitted, composed of a single layer 1.6 u thick. Micropyle absent. ‘Twenty- five sporulated oocysts measured 20-25 x 18-21 u, with a mean of 22.9 x 19.3 u. Their length-width ratios ranged from 1.0—1.3, with a mean of 1.19. Sporulated oocysts contain four ovoid sporocysts about 11.0—11.5 x 7-8 u; Stieda body present. Oocyst polar granule present. Oocyst residuum a homogeneous body. Sporocyst residual material composed of coarse granules. Sporozoites lie lengthwise in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Baiomys taylori (pigmy mouse). Other Host: Baiomys musculus (pigmy mouse). Location: Intestinal contents. Geographic Distribution: Mexico (Queretero, Oaxaca). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found by Levine, Ivens, and Kruidenier (1958) in one out of three B. taylori near Queretero, Queretero, Mexico, * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 65 and by Kruidenier, Levine, and Ivens (1960) in a B. musculus at Oaxaca, Mexico. EIMERIA ONYCHOMYSIS LEVINE, IVENS, AND KRUIDENIER, 1957 * (Plate 15, Fig. 104) Eimeria onychomysis Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts subspherical to ellipsoidal. Oocyst wall slightly rough, pale tan, composed of a single layer about 0.5 u thick. Micropyle absent. Six sporulated oocysts measured 20-21 x 17-20 u, with a mean of 20.4-18.6 u. Their length-width ratios ranged from 1.0-1.2, with a mean of 1.10. Sporulated oocysts with a polar granule and with one or several large, round, clear residual globules. Sporocysts about 11 x8 u, ovoid, with a Stieda body at one end. Sporocyst residuum formed of more or less compact coarse granules. Sporozoites lie with the large end of one beside the small end of the other. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Onychomys leucogaster (northern grasshopper mouse). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of two O. lewcogaster in Grand Canyon National Park, Arizona. EIMERIA PHYLLOTIS GONZALES-MUGABURU, 1942 Eimeria phyllotis Gonzales-Mugaburu, 1942: 137-151. Description: Oocysts elongate ellipsoidal, rose-colored. Micropyle ab- sent. Seven hundred sporulated oocysts from three host animals measured 22-30 x 12-16 u, with a mean of 26.2 14.0 u and a mean length-width ratio of 1.87. Oocyst wall 0.8-1.0 uw thick. Sporont 13-16 uw in diameter. Oocyst residuum and polar granule absent. Sporocysts 10-12 x 3-4 u, with granular residuum 3 u in diameter. Presence of Stieda body uncertain. Sporozoites 9-10 x 2-2.5 u. Sporulation Time: Two to three days in 3-5°% potassium dichromate. Schizogony and Gametogony: Macrogametes 18-20 uw in diameter. Microgametocytes 20-22 u in diameter. Microgametes 3 u long. The host cell nucleus is displaced. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 66 THE COCCIDIAN PARASITES OF RODENTS Prepatent Period: Five to eight days. Type Host: Phyllotis amicus amicus (?). Location: Epithelial cells of cecum. Geographic Distribution: South America (Peru). Pathogenicity: Unknown. Gonzales-Mugaburu (1942) found no gross lesions in infected rodents. Cross-Transmission Studies: Gonzales-Mugaburu (1942) was unable to infect young rats and white mice. Prevalence: Gonzales-Mugaburu (1942) found E. phyllotis in 38 per cent of 21 P. amicus from the Peru sierras. EIMERIA WEISSI GONZALES-MUGABURU, 1946 Eimeria weisst Gonzales-Mugaburu, 1946: 91-100. Description: Oocysts ovoid, reddish, with a double-membraned wall 1.2 u thick. Seven hundred oocysts measured 16-32 x 13-24 u, with a mean of 23.5 18.9 u. Their mean length-width ratio was 1.22. Micropyle absent. Oocyst polar granule apparently absent. Oocyst residuum present, 6-8 u in diameter, single at first and fragmenting later on. Sporocysts ellipsoidal, 10-11 x 6-7 u. Sporocyst residuum present. Presence of Stieda body unknown. Sporozoites lie longitudinally in sporocysts. Sporulation Time: Eight to nine days in 3 per cent potassium dichro- mate. Schizogony and Gametogony: Unknown. Prepatent Period: Four to six days. Type Host: Phyllotis amicus amicus (?). Location: Wleum and cecum. Geographic Distribution: South America (Peru). Pathogenicity: Unknown. Cross-Transmission Studies: Gonzales-Mugaburu (1946) was unable to transmit this species to white mice and guinea pigs. Prevalence: Gonzales-Mugaburu (1946) found this species in 9.5 per cent of 21 P. a. amicus from the Peruvian sierras. EIMERIA NEOTOMAE HENRY, 1932 Eimeria neotomae Henry, 1932a: 279-290. Description: Oocysts ellipsoidal. Oocyst wall smooth and transparent, apparently composed of a single layer. Henry (1932a) stated that the oocysts measured 16.0—22.4 x 12.8-19.2 u, with a mean of 22.4x 16.0 u, but one of the figures she gave for length must be wrong. A small micropyle is sometimes visible. Sporocysts subspherical, about 7.5 x 6.8 u, with an extremely thin wall which can be seen only with difficulty. Sporocysts just fill the oocyst without overlapping. Presence of oocyst GENUS Eimeria SCHNEIDER, 1875 67 polar granule uncertain; Henry (1932a) stated that there are usually no granules in the oocyst following sporulation, but that she saw a granule in an occasional oocyst. Oocyst residuum absent. Presence of sporocyst residuum unknown. Sporulation Time: Thirty-six to forty-eight hours in 2% potassium dichromate in small vials. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma fuscipes (wood rat). Location: Intestinal contents. Geographic Distribution: United States (California). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in 8 of 23 woodrats from the vicinity of Berkeley, California. EIMERIA RESIDUA HENRY, 1932 (Plate 35, Fig. 288) Eimeria residua Henry, 1932a: 279-290. Description: Oocysts subspherical. Oocyst wall comparatively thick, made up of two layers, the outer one brown and very rough, and the inner one clear and transparent. Oocysts 22-29 x 19-26 u, with a mean of 25.6 x 22.4 u. Micropyle absent. Sporulated oocysts with one or more polar granules and with a large residuum about 6 u in diameter com- posed of clear, homogeneous material. Sporocysts 107.5 u, pointed at one end, with a Stieda body. Presence of sporocyst residuum unknown. Sporulation Time: Eight to nine days in 2% potassium dichromate in small vials. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma fuscipes (woodrat). Location: Intestinal contents. Geographic Distribution: United States (California). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in 4 of 23 woodrats in the vicinity of Berkeley, California. EIMERIA OPERCULATA LEVINE, IVENS, AND KRUIDENIER, 1957 Eimeria operculata Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts ellipsoidal. Oocyst wall smooth, pale tan, about 1 u thick, composed of a single layer. Seven sporulated oocysts measured 68 THE COCCIDIAN PARASITES OF RODENTS 31-33 x 19-21 u, with a mean of 32.3 19.8 u. Their length-width ratios ranged from 1.6—-1.7, with a mean of 1.63. A circular suture around one end of the oocyst forms a cap or operculum about 7 u in diameter. There is no surface irregularity at the line of demarcation between the operculum and the oocyst proper, and the line itself is so fine that it can be seen only under high magnification. The pale suture line is visible between the operculum and the oocyst proper in high or low focus, but it is sometimes difficult to distinguish the line of demarcation or to recognize the operculum at all when focusing on the middle of the oocyst. Oocyst polar granule and oocyst residuum absent. Sporocysts sub- spherical, without Stieda body, about 7x6 u, containing two curled-up, banana-shaped sporozoites. Sporocyst residuum absent. Sporozoites re- leased from the sporocysts measure about 11x 2.2 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma stephensi (Stephens woodrat). Location: Intestinal contents. Geographic Distribution: United States (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of two N. stephensi in Grand Canyon National Park, Arizona. EIMERIA ALBIGULAE LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 15, Fig. 105) Eimeria albigulae Levine, Ivens, and Kruidenier, 1957: 80-88. Description: Oocysts spherical to subspherical. Oocyst wall composed of two layers: the outer layer colorless, rough, and about | u thick; the inner layer pale brownish, about 0.5 u thick. Forty-one sporulated oocysts measured 19-26 x 17-23 u, with a mean of 21.9 19.8 u. Their length- width ratios ranged from 1.0-1.4, with a mean of 1.10 and a standard deviation of 0.028. Micropyle absent. Sporulated oocysts contain one or two polar granules. Oocyst residuum composed of one to several large, clear globules. Sporocysts ovoid, approximately 11x9 uw with one end slightly pointed and with a slightly thickened wall at that end forming a small Stieda body. Sporozoites rounded, at ends of sporocyst. Sporocyst residuum represented by a number of more or less scattered granules. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma albigula (white-throated woodrat). Location: Intestinal contents. Geographic Distribution: United States (Arizona). GENUS Eimeria SCHNEIDER, 1875 69 Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one of two N. albigula in Grand Canyon National Park, Arizona. EIMERIA DAVISI IVENS, KRUIDENIER, AND LEVINE, 1959 (Plate 16, Fig. 106) Eimeria davisi Ivens, Kruidenier, and Levine, 1959: 53-57. Description: Oocysts subspherical to ellipsoidal. Oocyst wall composed of two layers: the outer one colorless to pale brownish yellow, smooth to slightly roughened, about 1.2—1.3 u thick; the inner one pale brownish to brownish, about 0.4—0.5 u thick. The layers can be separated by crush- ing the oocyst. Micropyle absent. Fifty-eight sporulated oocysts measured 22-32 x 21-24 u, with a mean of 27.6 x 22.8 u. Their length-width ratios ranged from 1|.1—1.4, with a mean of 1.21. Sporocysts ovoid; 23 sporocysts measured 10-12 x 7-9 u, with a mean of 11.1 8.3 u; their length-width ratios ranged from 1|.1—1.6, with a mean of 1.36. Oocyst polar granule present. Oocyst residuum composed of one to many clear globules up to 9 u or more in diameter; among 38 oocysts in which this feature was recorded, there was | globule in 30, 2 to 3 in 7, and many small ones in one oocyst. Sporocyst Stieda body medium-sized. Scattered residual granules usually present in sporocyst. Sporozoites lie lengthwise in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma albigula (white-throated wood rat). Location: Intestinal contents. Geographic Distribution: Mexico (Sonora). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in one out of five N. albigula near Agua Prieta, Sonora, Mexico. EIMERIA SP. BOYER AND SCORZA, 1957 Eimeria sp. Boyer and Scorza, 1957: 59-67. Description: Oocysts 21 x 13 u. Oocyst residuum very small. Sporozoites kidney-shaped. Sporocyst residuum present. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Zygodontomys brevicauda. Location: Intestine. Geographic Distribution: South America (Venezuela). 70 THE COCCIDIAN PARASITES OF RODENTS Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA SP. BOYER AND SCORZA, 1957 Eimeria sp. Boyer and Scorza, 1957: 59-67. Description: Oocysts 16x12 u. Oocyst residuum large. Sporozoites in the form of a closed C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Zygodontomys brevicauda. Location: Intestine. Geographic Distribution: South America (Venezuela). Pathogenicity: Unknown. Cross-Transmission Studies: None. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily CRICETINAE Host Tribe CRICETINI EIMERIA CRICETI NOLLER, 1920 Emend. PELLERDY, 1956 Eimeria falciformis var. criceti NOller, 1920: 180-182. Eimeria criceti Pellérdy, 1956: 75-102. Description: Odocysts spherical to subspherical, 11-22 u in diameter. Oocyst residuum absent. Sporocysts quite plump. Sporulation Time: Two to four days at room temperature. Schizogony and Gametogony: Noller (1920) did not describe these processes but stated that they were similar to those of E. falciformis of the mouse, except that they took place in the cecum and colon rather than in the small intestine. Prepatent Period: Unknown. Type Host: Cricetus cricetus (hamster). Location: Cecum and colon. The schizonts and gametocytes are found in the epithelial cells adjacent to the intestinal lumen, and not in those in the crypts. Geographic Distribution: Europe (Germany). Pathogenicity: Unknown. Hill (1952) reported that coccidiosis caused the death of a hamster in the London zoo, but did not describe the organism. Cross-Transmission Studies: None. Prevalence: This species was found in over 50 per cent of the young GENUS Eimeria SCHNEIDER, 1875 71 hamsters examined by Noller (1920) in Thtiringen. He never found it, however, in hamsters more than six weeks old, and animals found to be infected at two to four weeks of age were free of coccidia a few weeks later. Remarks: Although Noller (1920) described this species as a variety of E. falciformis whose host is Mus musculus, it occurs in a different part of the intestine, and in view of this and of the narrow host specificity of Eimeria species, Pellérdy (1956) rightly designated it Eimeria cricett. Since the specific name criceti was first used by Noller (1920) as a varietal designation, nomenclatorial practice retains him as the author. EIMERIA MIGRATORIA MUSAEV AND VEISOV, 1961 (Plate 16, Fig. 107) Eimeria migratoria Musaevy and Veisov, 1961: 971-975. Description: Oocysts spherical or subspherical. Oocyst wall smooth, colorless, composed of a single layer 1.5 uw thick. Micropyle absent. Fifty- six sporulated oocysts from four specimens were 15-23 uy in diameter with a mean of 19 u. Oocyst residuum absent. Two or sometimes three polar granules present. Sporocysts ovoid or spherical; ovoid sporocysts 6-11 x 4— 9 uw, with a mean of 107 u; spherical sporocysts 6-8 uw in diameter with a mean of 7 u. Stieda body absent. Sporocyst residuum composed of small granules located mostly between the sporozoites; a larger refractile body is also present between the sporozoites. Sporozoites bean-shaped. Sporulation Time: Two days at 25—-30C in 2.5°% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cricetulus migratorius (grey hamster). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961) found this species in 4 out of 64 specimens of Cricetulus migratorius from three regions of Nakhichevan ASSR, Azerbaidzhan SSR. EIMERIA IMMODULATA MUSAEV AND VEISOV, 1961 (Plate 17, Figs. 111 and 112) Eimeria immodulata Musaev and Veisov, 1961: 971-975. Description: Oocysts ovoid or ellipsoidal. Oocyst wall smooth, yellow- brown, composed of a single layer 1.0-1.5 uw thick. Micropyle absent. Fifty-two sporulated oocysts from five specimens were 18-24 12-19 u, 72 THE COCCIDIAN PARASITES OF RODENTS with a mean of 21x 16 u; their length-width ratios ranged from 1.2-1.6, with a mean of 1.4. Oocyst residuum absent. Oocyst polar granule some- times present. Sporocysts ovoid, 6-10 x 4-8 u, with a mean of 9x6 u. Stieda body absent. Sporocyst residuum composed of small granules located in different parts of the sporocyst. Sporozoites pear-shaped, rarely bean-shaped, with a prominent refractile globule in the broad end. Sporulation Time: Three days at 25-30C in 2.5% potassium dichro- mate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cricetulus migratorius (grey hamster). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961) found this species in 5 out of 64 speciments of C. migratorius from three localities in Nakhichevan ASSR, Azerbaidzhan SSR. EIMERIA CRICETULI MUSAEV AND VEISOV, 1961 (Plate 16, Fig. 108) Eimeria cricetuli Musaev and Veisov, 1961: 971-975. Description: Oocysts ovoid. Oocyst wall smooth, colorless, composed of a single layer 2 u thick. Micropyle absent. Twenty-five sporulated oocysts from one specimen measured 30-35 x 24-31 u, with a mean of 33 X 266.48 [szc] u; their length-width ratios ranged from 1.1-1.3, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid, 10-15 6-11 u, with a mean of 13x9 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites bean-shaped. Sporulation Time: Four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cricetulus migratorius (grey hamster). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961) found this species in 1 out of 64 C. migratorius from the Yardymlin region of Azerbaidzhan. GENUS Eimeria SCHNEIDER, 1875 73 EIMERIA JARDIMLINICA MUSAEV AND VEISOV, 1961 (Plate 16, Figs. 109 and 110) Eimeria jardimlinica Musaev and Veisov, 1961: 971-975. Description: Oocysts ovoid, rarely ellipsoidal. Oocyst wall smooth, colorless, composed of a single layer 1.5 uw thick. Micropyle absent. Four- teen sporulated oocysts from one specimen were 22-27 x 16-21 u, with a mean of 24x19 u; their length-width ratios ranged from 1.2-1.5, with a mean of 1.3. Oocyst residuum spherical or ovoid; ovoid residua 6-9 x 4—7 uw, with a mean of 8x6 u; spherical residua 6-8 u in diameter, with a mean of 7 u. Oocyst polar granule absent. Sporocysts ovoid, 10-13 x 6-9 u, with a mean of 117 u. Stieda body absent. Sporocyst residuum composed of small granules often located between the sporozoites. Sporozoites comma-shaped, with a refractile globule in the broad end. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cricetulus migratorius (grey hamster). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961) found this species in | out of 64 C. migratorius from the Yardymlin region of Azerbaidzhan. EIMERIA ARUSICA MUSAEV AND VEISOV, 1961 (Plate W7, Fig. 113) Eimeria arusica Musaev and Veisov, 1961: 971-975. Description: Oocysts spherical or subspherical, yellow. Oocyst wall smooth, composed of two layers each | u thick, the outer layer colorless and the inner layer dark brown. Micropyle absent. ‘Twenty-six sporulated oocysts from two specimens were 20-26 u in diameter, with a mean of 23 wu. Oocyst residuum spherical or ovoid (illustrated as a homogeneous body); spherical residua 6-8 uw in diameter, with a mean of 7 u; ovoid residua 8X6 u. Oocyst polar granule absent. Sporocysts ovoid, 10-13 x 6-9 u, with a mean of 11 x7 u. Stieda body present. Sporocyst residuum composed of small granules located between the sporozoites. Sporozoites comma- or bean-shaped. Sporulation Time: Four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. 74 THE COCCIDIAN PARASITES OF RODENTS Prepatent Period: Unknown. Type Host: Cricetulus migratorius (grey hamster). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961) found this species in 2 out of 64 C. migratorius from the outskirts of the village of Arus in the Yardymlin region of Azerbaidzhan. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily MICROTINAE Host Tribe LEMMINI EIMERIA DICROSTONICIS LEVINE, 1952 (Blate Wy Figs 120) Eimeria dicrostonicis Levine, 1952 (1951): 205-208. Description: Oocysts ellipsoidal. Oocyst wall composed of two layers: the outer one yellowish brown, a little more than 1| u thick, with a rough, pitted surface; the inner layer colorless and about 0.5 u thick. Micropyle absent. Twenty-nine oocysts from two lemmings measured 23-27 x 27-31 u, with a mean of 24.8x 29.1 u. Their length-width ratios ranged from 1.1-1.3, with a mean of 1.2. One or occasionally two oocyst polar granules present. Oocyst residuum absent, occasionally represented by a few small granules. Sporocysts 7-9 x 13-15 u, with a mean of 8.4 x 14.0 u; sporocyst length-width ratio 1.5-1.8, with a mean of 1.7. Sporocysts ellipsoidal but slightly pointed at either end. Either no Stieda body or a very small one present. Sporocyst residuum absent, occasionally represented by a few small granules. Sporozoites lie lengthwise in the sporocysts, with the broad end of one beside the narrow end of the other. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dicrostonyx groenlandicus richardsoni (varying lemming). Location: Feces. Geographic Distribution: North America (Canadian Northwest Ter- ritory). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine (1952) found this species in 5 of 14 D. g. richardsoni from the mouth of the Perry River on the Arctic Ocean. sI oO GENUS Eimeria SCHNEIDER, 1875 Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily MICROTINAE Host Tribe MICROTINI EIMERIA ARVICOLAE (GALLI-VALERIO, 1905) REICHENOW, 1921 Coccidium arvicolae Galli-Valerio, 1905: 519-522. Eimeria arvicolae (Galli-Valerio) Reichenow, 1921: 1136-1277; Iwanoff-Gobzem, 1934: 149-151; Svanbaevy, 1956: 180-191; Svanbaev, 1958: 183-186 (?). Eimeria arvalis Iwanoft-Gobzem, 1934: 149-151 (an invalid emendation, with no nomenclatural status). Eimeria musculi Yakimoff and Gousseff. Svanbaev, 1956: 180-191 (from Microtus arvalis). [non] Eimeria musculi Yakimoff and Gousseff, 1938: 1-3; Svanbaev, 1956: 180-191 (from Apodemus sylvaticus—see E. russiensis) (from Mus musculus—see E. muscullt). [non] Eimeria arvicolae: Musaev and Veisov, 1960: 51-61 (see E. batabatensis). Description: Oocysts spherical, 14-18 u in diameter. Oocyst wall clear, with double contour. Sporont cytoplasm finely granular, somewhat yel- lowish. The above is all the information given by Galli-Valerio (1905) in his original description of this species from Microtus nivalis. The form described by Iwanoff-Gobzem (1934) under this name from M. arvalis had ovoid oocysts, 13-28 x 11-21 u, with a mean of 20.1 x 16.3 u. Their length-width ratios ranged from 1.1—1.2, with a mean of 1.18. They lacked oocyst and sporocyst residua. No other information was given. Svanbaev (1956) described two forms from M. arvalis. One form, designated E. arvicolae, had ovoid, greenish oocysts with a smooth, double-contoured wall 1.5 uw thick and without a micropyle. Its oocysts measured 22.5 18.8 uw. Oocyst polar granule and oocyst and sporocyst residua were absent. The sporocysts were ovoid and 12.3 x 8.8 u. Svan- baev’s second form, which he designated E. musculi, differed from the first form only in having spherical oocysts 22.0 u in diameter, sporocysts 13.2 x 8.8 u, and somewhat smaller sporozoites. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Microtus (syn., Arvicola) nivalis (snow vole). Other Hosts: Microtus arvalis (common European vole); Alticola strelzovi (Strelzov’s vole) (?). Location: Intestine. Geographic Distribution: Europe (Switzerland), USSR (Kazakhstan). Pathogenicity: Unknown. 76 THE COCCIDIAN PARASITES OF RODENTS Cross-Transmission Studies: None. Prevalence: Iwanoft-Gobzem (1934) found her form in 9 out of 22 M. arvalis in Kazakhstan. Svanbaev (1956) found both his spherical and ovoid forms in 2 out of 33 M. arvalis in western Kazakhstan; they were present in the same individuals. Remarks: Although Iwanoff-Gobzem (1934) believed that the form she found in Microtus arvalis was identical with that described by Galli- Valerio (1950) from M. nivalis, it differed from it in being ovoid rather than spherical. However, both of their descriptions were very incomplete. Svanbaev (1956) found what he considered two species of Eimeria in M. arvalis. The only essential difference between them was in shape. He considered the ovoid form to be E. arvicolae and the spherical one E. musculi. However, if Galli-Valerio’s (1905) original description had been followed, Svanbaev should have referred the spherical form to E. arvicolae. Furthermore, in view of the failure of all attempts at trans- mitting Eimeria from one rodent genus to another, it is highly unlikely that the same species would occur in two rodents of different subfamilies. E. musculi was originally described from Mus musculus (subfamily Murinae), while Microtus belongs to the subfamily Microtinae. Finally, the great similarity between Svanbaev’s two forms from M. arvalis and the fact that they were both found in the same two voles out of 33 examined suggest that they most likely belong to the same species. Svanbaev (1958) reported finding FE. arvicolae oocysts in 4 of 43 Strelzov’s voles (Alticola strelzovi) in Central Kazakhstan. He did not describe them. EIMERIA MICROTINA MUSAEV AND VEISOV, 1959 (Plate 17, Fig. 114) Eimeria microtina Musaev and Veisov, 1959: 45-50. Description: Oocysts spherical, sometime slightly ovoid. Oocyst wall smooth, colorless, 0.8 u thick, apparently composed of a single layer. Micropyle absent. Fifty sporulated oocysts from one animal were meas- ured. The spherical ones were 16 uw in diameter and the ovoid ones were 12-18 x 11-16 uw, with a mean of 16X15 wu. Oocyst residuum present. Oocyst polar granule absent. Sporocysts ovoid, 5-8 x4—5 u, with a mean of 74 u. Stieda body conspicuous. Sporocyst residuum present, com- posed of rather small granules. Sporozoites illustrated with a clear globule at the large end. Sporulation Time: Three days in 2.5°% potassium dichromate solution at 25-30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. GENUS Eimeria SCHNEIDER, 1875 Wa Sporogony: Not described. Type Host: Microtus socialis (social vole). Location: Intestinal contents. Geographic Distribution: USSR. Musaev and Veisov (1959) found the host animal in a freight car carrying goods from Iran to Dzhul’fa, Azerbaidzhan. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA DZHULFAENSIS MUSAEV AND VEISOV, 1959 Eimeria dzhulfaensis Musaev and Veisov, 1959: 45-50. Description: Oocysts spherical, sometimes slightly ovoid. Oocyst wall rough, composed of two layers, of which the outer is 1.2 u thick and dark brown and the inner is 0.2 u thick. Micropyle absent. Fifty sporu- lated oocysts were measured. The spherical ones were 24 u in diameter and the ovoid ones were 21—25~x 20-24 u, with a mean of 24x21 u. Oocyst residuum and polar granule present. Sporocysts ovoid with a pointed end, 11-14x6-9 u, with a mean of 12.5x8 u. Stieda body prominent. Sporocyst residuum present, composed of fine granules. Sporozoites pear-shaped, with a small globule at the broad end. Sporulation Time: Three days in 2.5% potassium dichromate solution at 25-30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: Not described. Type Host: Microtus socialis (social vole). Location: Intestinal contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1959) found this species in 11 (6 per cent) of 169 voles from Dzhul’fin and other regions of Azerbaidzhan. EIMERIA WENRICHI SAXE, LEVINE, AND IVENS, 1960 (Plate 17, Figs. 115 and 116) Eimeria wenrichi Saxe, Levine, and Ivens, 1960: 61-63. Eimeria species A. Saxe, 1952: 13. Eimeria species B. Saxe, 1952: Ibid. Description: Oocysts ellipsoidal to ovoid. Oocyst wall composed of a single layer. Micropyle absent. Oocysts of two nonoverlapping sizes but without any other morphological differences. Fifty-six of the larger 78 THE COCCIDIAN PARASITES OF RODENTS oocysts measured 16-22 x 12-16 u, with a mean of 18.9x 14.3 u; their length-width ratios ranged from 1.1—1.6, with a mean of 1.32. Fourteen sporocysts of this form measured 9-11 x 5-8 u, with a mean of 9.7 x 6.0 u; their length-width ratios ranged from 1.3-2.0, with a mean of 1.62. Thirty-one oocysts of the smaller form measured 11-15 x 8-11 u, with a mean of 12.8 x 9.8 u; their length-width ratios ranged from 1.1—1.7, with a mean of 1.31. Ten sporocysts of this form measured 6.5—-7.5 4 u, with a mean of 6.94.0 u; their length-width ratios ranged from 1.6-1.9, with a mean of 1.72. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, with Stieda body. Sporocyst residuum present, compact. Sporulation Time: Two to three days at room temperature (24—27C) in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Microtus pennsylvanicus (meadow mouse). Location: Cecal contents. Geographic Distribution: United States (Pennsylvania). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in a single meadow mouse. EIMERIA BOHEMIGA RYSAVY, 1957 (Plate 17, Figs. 117 and 118) Eimeria bohemica RySavy, 1957: 331-336. Description: Oocysts ellipsoidal, pale yellowish brown. Oocyst wall ap- parently smooth, illustrated as composed of a single layer. Micropyle prominent, 4—4.4 yw wide. Oocysts 19-23x11-13 u, with a mean of 20 x 12 u; most oocysts measured 21 x 11 u. Oocyst residuum a small, dark, spherical body 2 u in diameter. Oocyst polar granule apparently absent. Sporocysts ovoid, 9.56 u. Sporocyst residuum present, finely granular. Stieda body not illustrated. Sporulation Time: Three days in 2% potassium dichromate solution at 24C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicola terrestris (water vole). Location: Feces. Geographic Distribution: Europe (Czechoslovakia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: RySavy (1957) found this species in 3 out of 21 A. ter- restris near Jindiichova Hradce, southern Czechoslovakia. GENUS Eimeria SCHNEIDER, 1875 79 EIMERIA BATABATENSIS N. SP. (Plate 17, Fig. 119) Eimeria arvicolae Galli-Valerio. Musaev and Veisov, 1960: 51-61. Description: Oocysts ovoid, colorless. Oocyst wall smooth, described as double-contoured but illustrated with a single layer, | u thick. Micropyle absent. Forty-one sporulated oocysts measured 14-20 x 10-16 u, with a mean of 19x15 u; their length-width ratios ranged from 1.1-1.5, with a mean of 1.32. Oocyst residuum and polar granule absent. Sporocyst ovoid, rarely spherical; ovoid sporocysts 6-8 x 4-6 u, with a mean of 7x4 wu; spherical sporocysts 4-6 uw in diameter, with a mean of 6 u. Sporocyst residuum present, composed of small granules. Stieda body absent. Sporozoites comma-shaped, with a small globule at the large end. Sporulation Time: Two days in 2.5% potassium dichromate at 25—30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicola terrestris (water vole). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Musaev and Veisov (1960) found this species in water voles on the summer pastures of Batabat (2400 m above sea level) in the Shakhbuz region of Nakhichevan, Azerbaidzhan SSR. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. Remarks: Although Musaev and Veisov (1960) called this form Eimeria arvicolae, it differs morphologically from that species in having a sporocyst residuum. In addition, it occurs in a different host genus. (The hosts of E. arvicolae are Microtus nivalis and M. arvalis; both of these voles were formerly thought to be members of the genus Arvicola; however, according to Ellerman and Morrison-Scott (1951), the only species now accepted in this genus is A. terrestris.) We are therefore naming it Eimeria batabatensis n. sp. EIMERIA TERRESTRIS MUSAEV AND VEISOV, 1960 (Plate 18, Fig. 121) Eimeria terrestris Musaev and Veisov, 1960: 51-61. Descriptions: Oocysts ovoid, rarely ellipsoidal, colorless or light yellow. Oocyst wall smooth, tri-contoured and double-layered, 1.5-2 u thick; outer layer colorless to yellowish, inner layer dark yellow or brown. Micropyle prominent. Fifty sporulated oocysts from six animals measured 18-22 x 12-16 u, with a mean of 21 x 16 u; their length-width ratios ranged from 1.2-1.5, with a mean of 1.37. Oocyst residuum and polar granule 80 THE COCCIDIAN PARASITES OF RODENTS absent. Sporocysts usually ovoid, rarely spherical, 6-8 x 4-6 u, with a mean of 8x6 u. Sporocyst residuum present, composed of small granules which often form a ball. Stieda body absent. Sporozoites pear-shaped, rarely comma-shaped. Sporulation Time: Three to 3.5 days in 2.5% potassium dichromate at 25 to 30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicola terrestris (water vole). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Musaev and Veisov (1960) found this species in water voles on the summer pastures of Batabat (2400 m. above sea level) in the Shakhbuz region of Nakhichevan, Azerbaidzhan SSR. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA TALISCHAENSIS MUSAEV AND VEISOV, 1960 (Plate 18, Fig. 122) Eimeria talischaensis Musaev and Veisov, 1960: 51-61. Description: Oocysts spherical, colorless. Oocyst wall smooth, tri-con- toured and double-layered, 1.5—2 u thick; each layer | u thick; outer layer colorless, inner layer light yellowish brown. Micropyle absent. Ten sporulated oocysts were 18-22 u in diameter, with a mean of 21 u. Oocyst residuum absent. Oocyst polar granule present. Sporocysts spheri- cal or ovoid; spherical sporocysts 4—8 u in diameter, with a mean of 6 u; ovoid sporocysts 4—8 x 2-8 yu, with a mean of 6 x 5.6 u. Sporocyst residuum composed of small granules. Stieda body absent. Sporozoites comma- shaped, rarely pear-shaped, with a clear globule at the broad end. Sporulation Time: Two days in 2.5% potassium dichromate at 25—30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicola terrestris (water vole). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Musaev and Veisov (1960) found this species in water voles on the summer pastures of Batabat (2400 m. above sea level) in the Shakhbuz region of Nakhichevan and in the mountain zone of the Yadymlin region, both in Azerbaidzhan. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. GENUS Eimeria SCHNEIDER, 1875 81 EIMERIA SP. (RYSAVY, 1954) Eimeria falciformis (Eimer). RySavy, 1954: 131-174 (pro parte). [non] Eimeria falciformis (Eimer, 1870) Schneider, 1875: xl-xlv. Description: Uncertain. Host: Microtus arvalis (vole). Location: Feces. Geographic Distribution: Czechoslovakia. Remarks: RySavy (1954) reported E. falciformis from Mus musculus, Apodemus flavicollis, A. sylvaticus, Microtus arvalis, and Clethrionomys glareolus in Czechoslovakia. The description he gave was similar to that of E. falciformis, but he did not describe the forms from each host separately. He was apparently unaware of Galli-Valerio’s (1932, 1940) and Pellérdy’s (1954) descriptions of six species of Eimeria from Apodemus or of Iwanofl-Gobzem’s (1934) report of E. arvicolae from Microtus arvalis, nor was he aware of Pellérdy’s (1954) inability to trans- mit any of his Apodemus coccidia to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, or Cricetus cricetus. It is very doubtful that the forms RySavy saw in the hosts other than M. musculus were E. falciformis. However, since he did not describe them separately from each host, it is useless to attempt to assign names to them. EIMERIA ONDATRAZIBETHICAE MARTIN, 1930 Emend. Plate 35, Figs. 289 and 290) Eimeria ondatrae-zibethicae Martin, 1930: 273-278; Svanbaev, 1962a: 206-207. Eimeria stiedae (Lindemann) Kisskalt and Hartmann. Law and Kennedy, 1932. [non] Eimeria stiedae (Lindemann, 1865) Kisskalt and Hartmann, 1907. Description: Oocysts mostly ellipsoidal, but also spherical, ovoid, or cylindrical. Seventy-five oocysts measured 19-28 x 13-26 u, with a mean of 22.3 x 18.0 u. One spherical oocyst was 10 u in diameter. Oocyst length- width ratio 1.0-1.7, with a mean of 1.14. Oocyst wall brownish, very thick, smooth, with a double contour, occasionally radially striated, sometimes with one end flattened. Micropyle well developed in some oocysts and not visible in others. Sporocysts 12-17 x 8-11 u, with a mean of 13.9x9.7 u. A small oocyst residuum may or may not be present. Oocyst polar granule not mentioned or illustrated. Sporocysts with Stieda body and small residuum. According to Svanbaev (1962a), the oocysts are ellipsoidal or spherical, 22-35 x 18-27 u, with a mean of 28.5 22.5 u; their length-width ratio ranges from 1.2—-1.3, with a mean of 1.3; the oocyst wall is smooth, greenish to yellow-green, 1.3-1.8 u thick, with a micropyle; an oocyst polar granule is absent; the sporocysts are spherical, ellipsoidal, or ovoid, 13-15x8-9 u, with a mean of 14x8.5 u; the sporocyst residuum is in the form of small granules; the sporozoites are comma-shaped or piriform, 8-11 x 3-5 u, with a mean of 9.5 4 u. 82 THE COCCIDIAN PARASITES OF RODENTS Sporulation Time: Five days in 5% potassium dichromate at 21—27C. Schizogony and Gametogony: One microgametocyte measured by Martin (1930) was 10x 11 u and another was 22 x 23 u. One macrogamete was 15x17 u and another was 15x30 u. These were apparently in epithelial cells of the jejunum. Prepatent Period: Unknown. Type Host: Ondatra zibethica (muskrat). Location: Jejunum, liver. Geographic Distribution: North America (Nebraska, Maryland, New York, Wisconsin, Ontario); USSR (Kazakhstan). Pathogenicity: This species can be highly pathogenic. In an outbreak described by Martin (1930) among muskrats kept under a condition of semi-domestication in Nebraska, about 100 animals died in a com- paratively short period. Deaths occurred most commonly among the young animals, and whole litters sometimes succumbed. Shillinger (1938) reported high mortality among wild muskrats in Wisconsin during a drought when the water level was low and the animals’ passageways became muddy and contaminated with feces. LeCompte (1933) reported a death due to coccidiosis in a muskrat in Maryland. Martin (1930) described the lesions caused by this species. He found an intense hemorrhagic enteritis of the jejunum. The intestinal lumen was filled with blood and debris. Microscopically, the intestinal epithe- lium was desquamated and in some instances almost completely denuded. There was extensive hemorrhage into the lumen of the intestine. Vascular congestion and round cell infiltration were noted. The liver contained many small, white foci up to 2 mm. in diameter. They were due to focal necrosis and vascular congestion. Oocysts were also found in the liver. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962a) found this coccidum in 21 per cent of 38 O. zibethica in northern Kazakhstan. Prevalence: Bump (1942) found Eimeria sp. (presumably E. ondatra- zibethicae) in 31 of 91 muskrats in New York. He did not describe them. EIMERIA RYSAVYI N. SP. (Plate 18, Figs. 123, 124, and 125) Eimeria apodemi Pellérdy. RySavy, 1957: 331-336. [non] Eimeria apodemi Pellérdy, 1954: 187-191. Eimeria hindlei Yakimoff and Gousseff. RySavy, 1954: 131-174 (pro parte); Cerna, 1962: 1-13. [non] Eimeria hindlei Yakimoft and Gousseff, 1938: 1-3. Description: According to Rysavy (1957), the oocysts are broadly el- lipsoidal, a few almost subspherical, and some rather asymmetrical, 23-29 x 18-23 u, with a mean of 25x20 u; most of the oocysts were GENUS Eimeria SCHNEIDER, 1875 83 25x19 u. Oocyst wall rather thick, pale yellowish brown, illustrated as smooth and composed of a single layer. Micropyle absent. Sporocysts ovoid. Sporocyst residuum large and prominent. RySavy (1957) gave no other morphological information; he illustrated an unsporulated oocyst. According to Gerna (1962), the oocysts were mostly ellipsoidal (‘‘oval”’), sometimes ovoid, 20-30 x 17-21 u; the oocyst wall was smooth, illustrated as composed of a single layer, yellowish to light brown; a micropyle was absent. The sporocysts were ellipsoidal (“oval”), 11-12 x 6-7 u, illustrated without a Stieda body. Oocyst polar granule and oocyst residuum were absent, but a small amount of sporocyst residual material was present. The sporozoites lay lengthwise in the sporocysts; each contained a clear globule at one end. Sporulation Time: Four to five days in 2% potassium dichromate solution. Schizogony: Unknown. Gametogony: According to Gerna (1962), gametogony occurs primarily in the duodenum and jejunum, and only occasionally in the ileum. The sexual stages are found in the mucosa and submucosa. The mature microgametocytes measured 29-44 x 18-33 u. The microgametes were about 2 u long. The mature macrogametes measured 16-23 x 17.5 u. Prepatent Period: Unknown. Type Host: Clethrionomys glareolus (common red-backed vole). Location: Sexual stages in duodenum, jejunum, and occasionally ileum. Geographic Distribution: Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Ry’avy (1957) found this species in 2 out of 14 C. glareolus from the vicinity of Jindtichova Hradce, southern Czechoslovakia. Cerna (1962) found it in Mustek, Sumava, Czechoslovakia. Remarks: RySavy (1957) reported this species in the red-backed vole under the name Eimeria apodemi. Since, however, Pellérdy (1954) was unable to transmit E. apodemi from Apodemus to Clethrionomys, this identification cannot be correct, and we are naming RySavy’s form Eimeria rysavyi n. sp. Rysavy (1954) also described what was undoubtedly the same form under the name Eimeria hindlei from C. glareolus and Apodemus sylvaticus in Czechoslovakia. He did not differentiate between the forms from the two hosts, the oocysts failed to sporulate, and he was apparently unaware at that time of Galli-Valerio’s (1932, 1940) and Pellérdy’s (1954) descriptions of six species of Eimeria from Apodemus or of Pellérdy’s inability to transmit any of his Apodemus coccidia to Mus musculus or C. glareolus. Since the type host of E. hindlei is Mus musculus, it is clear that RySavy’s form does not belong to this species. 84 THE COCCIDIAN PARASITES OF RODENTS Cerna (1962) gave further information regarding what was undoubtedly the same species from C. glareolus, referring it to RySavy’s “E. hindlei” from this host. EIMERIA CERNAE N. SP. (Plate 19, Fig. 132) Eimeria schueffneri Yakimoft and Gousseff. Cerna, 1962: 1-13. [non] Eimeria schueffnert Yakimoff and Gousseff, 1938: 1-3. Description: Oocysts 13-23 x 11-17 u, ellipsoidal, with a very thin wall. Micropyle absent. Oocyst residuum absent. Oocyst polar granule some- times present. Sporocysts mostly ellipsoidal (“oval”), 9-15x4-7 u; oc- casionally spherical, 6 u. Sporocyst residuum and Stieda body absent. Sporozoites illustrated as lying longitudinally in sporocysts. Sporulation Time: Two to four days in 1.5% potassium dichromate solution. Schizogony: Unknown. Gametogony: According to Cerna (1962), gametogony occurs in the epithelium of the cecum. The mature microgametocytes are 8-15 x 6-11 u and the mature macrogametes 11-15 x 6-12 u. Prepatent Period: Unknown. Type Host: Clethrionomys glareolus (common red-backed vole). Location: Cecum. Geographic Distribution: Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. Remarks: Cerna (1962) described this species under the name E. schueffnert. However, the latter is a parasite of Mus musculus, and, for the reasons given under E. rysavyi, this identification cannot be correct; therefore, we are naming Cerna’s form Eimeria cernae n. sp. Cerna (1962) said in her text that she found the same form in Apodemus sylvaticus, and in her table she listed it in Microtus arvalis and “P. subterraneus.” However, in the absence of any description of the forms from these hosts, it is impossible to know what species she was dealing with. EIMERIA SP. (RYSAVY, 1954) Eimeria falciformis (Eimer). Ry$avy, 1954: 131-174 (pro parte); Cerna and Daniel, 1956: 19-23 (pro parte). [mon] Eimeria falciformis: Schneider, 1875: xl-xlv. Description: Uncertain. Host: Clethrionomys glareolus (common red-backed vole). GENUS Eimeria SCHNEIDER, 1875 85 Location: Feces. Geographic Distribution: Czechoslovakia. Remarks: RySavy (1954) reported E. falciformis from Mus musculus, Apodemus flavicollis, A. sylvaticus, Microtus arvalis, and Clethrionomys glareolus in Czechoslovakia. For the reasons given in the discussion of these forms in Microtus arvalis (p. 81) it is very doubtful that the forms RySavy saw in the hosts other than M. musculus were E. falciformis, but it is useless to attempt to assign names to them. Cerna and Daniel (1956) reported E. falciformis from Clethrionomys glareolus and Apodemus flavicollis in Czechoslovakia. They gave the same dimensions as RySavy, but mentioned that the oocyst wall was more than 1 u thick. For the same reasons, these forms were undoubtedly not E. falciformis, but cannot be assigned separate names. EIMERIA (?) SP. CERNA and DANIEL, 1956 Eimeria sp. Cerna and Daniel, 1956: 19-23. This form, which failed to sporulate, was found by Cerna and Daniel (1956) in Clethrionomys glareolus and Apodemus flavicollis in Czecho- slovakia. For further information see its description under 4. flavicollis (onal ls) Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily MICROTINAE Host Tribe ELLOBIINI EIMERIA ELLOBIT SVANBAEV, 1956 (Plate 36, Fig. 293) Eimeria ellobti Svanbaev, 1956: 180-191. Description: Oocysts ovoid, greenish, 30.1 x 24.2 u, with a smooth wall 1.3-1.7 u thick composed of two layers. Oocyst length-width ratio 1.25. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts spherical, 8.8 u in diameter. Stieda body presumably absent. Sporocyst residuum present. Sporozoites kidney-shaped, 6.55.6 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Ellobius talpinus (mole lemming). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. 86 THE COCCIDIAN PARASITES OF RODENTS Prevalence: Svanbaev (1956) found this species in three out of ten E. talpinus in western Kazakhstan. ‘The same three individuals were infected with Eimeria talpini and E. kazakhstanensis. EIMERIA KAZAKHSTANENSIS N. SP. Eimeria volgensis Sassuchin and Rauschenbach. Svanbaev, 1956: 180-191. [non] Eimeria volgensis Sassuchin and Rauschenbach, 1932: 646-650. Description: Oocysts ovoid, greenish to occasionally yellowish brown, with a smooth, colorless [sic], single-layered wall 1.6 uw thick. Oocysts 27.5 x 24.2 uw. Oocyst length-width ratio 1.14. Micropyle at narrow end of oocyst, small. Oocyst polar granule present in unsporulated oocyst. Oocyst residuum absent. Sporocysts ovoid or pear-shaped, 11-13 x 7-8 u, with a mean of 12.2 x 7.2 u. Stieda body not mentioned. Sporocyst residual granules present. Sporozoites oval, 8.45.9 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Ellobius talpinus (mole lemming). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaey (1956) found this species in three out of ten E. talpinus in western Kazakhstan. The same three individuals were infected with Eimeria ellobu and E. talpini. Remarks: Svanbaev (1956) described this species under the name E. volgensis Sassuchin and Rauschenbach, 1932. He did not give an illustra- tion. While it was similar in general to this species, it had a small rather than a large micropyle and apparently did not have the pointed micro- pylar end characteristic of E. volgensis. Furthermore, the two hosts be- long to widely separate rodent groups. Spermophilus, the host genus of E. volgensis, is a member of the suborder Sciuromorpha, while Ellobius is a member of the suborder Myomorpha. Cross-transmission of Ezmeria between these genera is extremely doubtful, and in the absence of proof to the contrary should be ruled out. Hence we have assigned this form the name Eimeria kazakhstanensis n. sp. EIMERIA TALPINI N. SP. Eimeria beckeri Yakimoff and Sokoloff. Svanbaev, 1956: 184-185 (in Ellobius talpinus). Description: Oocysts spherical, 24.2 u in diameter. Oocyst wall smooth, colorless, 1.5 uw thick, composed of two layers. Micropyle absent. Oocyst polar granule absent. Oocyst residuum absent. Sporocysts oval, 11.0 x 8.8 u. Stieda body not mentioned. Sporocyst residual granules present. GENUS Eimeria SCHNEIDER, 1875 87 Sporozoites 6.53.7 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Ellobius talpinus (mole lemming). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in three out of ten E. talpinus in western Kazakhstan. The same three individuals were infected with Eimeria ellobii and E. kazakhstanensis. Remarks: Svanbaev (1956) described this species under the name E. beckeri Yakimoff and Sokoloff. He did not give an illustration. While it was similar in general to this species, it was always spherical rather than ovoid to spherical. Furthermore, the two hosts belong to widely separate rodent groups. Spermophilus, the host genus of E. beckeri, is a member of the suborder Sciuromorpha, while Ellobius is a member of the suborder Myomorpha. For the reasons given under E. kazakhstanensis (p. 86), we are assigning this form the name E. talpini n. sp. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily GERBILLINAE EIMERIA MERIONIS MACHUL’SKH, 1949 Eimeria merionis Machul’skii, 1949. Eimeria marionis [sic]: Svanbaev, 1956: 180-191; Svanbaev, 1962: 23-39. Description: Original form from Meriones unguiculatus: Oocysts ovoid, 17-23 x 15-19 u, with a mean of 19.1 x 15.3 u. Oocyst wall 1.0-1.2 u thick, double-contoured, composed of a single layer, orange. Micropyle not mentioned. Oocyst polar granule present. Oocyst and sporocyst residua present. Sporocysts 8-106 u. Form from M. tamariscinus: The following description is from Svan- baev (1962): Oocysts ellipsoidal or subspherical, 19-21 x 17-18 u, with a mean of 21 x 17.5 u; length-width ratio 1.2. Oocyst wall smooth, yellow- orange or yellow-brown, 0.9-1.3 u thick. Micropyle absent. Oocyst polar granule usually present. Oocyst residuum a large, granular sphere be- tween the sporoblasts. Sporocysts ellipsoidal or subspherical, 9-13 x 6-8 u, with a mean of 11 x7 u. Sporozoites comma-shaped, 5-7 x 2-4 u, with a mean of 6x3 u. Sporocyst residuum in form of small granules. Schizogony and Gametogony: Presumably unknown. Prepatent Period: Presumably unknown. 86 THE COCCIDIAN PARASITES OF RODENTS Prevalence: Svanbaev (1956) found this species in three out of ten E. talpinus in western Kazakhstan. The same three individuals were infected with Eimeria talpini and E. kazakhstanensis. EIMERIA KAZAKHSTANENSIS N. SP. Eimeria volgensis Sassuchin and Rauschenbach. Svanbaev, 1956: 180-191. [non] Eimeria volgensis Sassuchin and Rauschenbach, 1932: 646-650. Description: Oocysts ovoid, greenish to occasionally yellowish brown, with a smooth, colorless [sic], single-layered wall 1.6 uw thick. Oocysts 27.5 x 24.2 uw. Oocyst length-width ratio 1.14. Micropyle at narrow end of oocyst, small. Oocyst polar granule present in unsporulated oocyst. Oocyst residuum absent. Sporocysts ovoid or pear-shaped, 11-13 x 7-8 u, with a mean of 12.2 x 7.2 u. Stieda body not mentioned. Sporocyst residual granules present. Sporozoites oval, 8.45.9 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Ellobius talpinus (mole lemming). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in three out of ten E. talpinus in western Kazakhstan. The same three individuals were infected with Eimeria ellobu and E. talpini. Remarks: Svanbaev (1956) described this species under the name E. volgensis Sassuchin and Rauschenbach, 1932. He did not give an illustra- tion. While it was similar in general to this species, it had a small rather than a large micropyle and apparently did not have the pointed micro- pylar end characteristic of E. volgensis. Furthermore, the two hosts be- long to widely separate rodent groups. Spermophilus, the host genus of E. volgensis, is a member of the suborder Sciuromorpha, while Ellobius is a member of the suborder Myomorpha. Cross-transmission of Ezmeria between these genera is extremely doubtful, and in the absence of proof to the contrary should be ruled out. Hence we have assigned this form the name Eimeria kazakhstanensis n. sp. EIMERIA TALPINI N. SP. Eimeria beckeri Yakimoft and Sokoloff. Svanbaev, 1956: 184-185 (in Ellobius talpinus). Description: Oocysts spherical, 24.2 u in diameter. Oocyst wall smooth, colorless, 1.5 u thick, composed of two layers. Micropyle absent. Oocyst polar granule absent. Oocyst residuum absent. Sporocysts oval, 11.0 x 8.8 u. Stieda body not mentioned. Sporocyst residual granules present. GENUS Eimeria SCHNEIDER, 1875 87 Sporozoites 6.53.7 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Ellobius talpinus (mole lemming). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in three out of ten E. talpinus in western Kazakhstan. The same three individuals were infected with Eimeria ellobi and E. kazakhstanensis. Remarks: Svanbaev (1956) described this species under the name E. beckeri Yakimoff and Sokoloff. He did not give an illustration. While it was similar in general to this species, it was always spherical rather than ovoid to spherical. Furthermore, the two hosts belong to widely separate rodent groups. Spermophilus, the host genus of EF. beckeri, is a member of the suborder Sciuromorpha, while Ellobius is a member of the suborder Myomorpha. For the reasons given under EF. kazakhstanensis (p. 86), we are assigning this form the name E. talpini n. sp. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily GERBILLINAE EIMERIA MERIONIS MACHUL’SKII, 1949 Eimeria merionis Machul’skii, 1949. Eimeria marionis [sic]: Svanbaev, 1956: 180-191; Svanbaev, 1962: 23-39. Description: Original form from Meriones unguiculatus: Oocysts ovoid, 17-23 x 15-19 u, with a mean of 19.1 x 15.3 uw. Oocyst wall 1.0-1.2 u thick, double-contoured, composed of a single layer, orange. Micropyle not mentioned. Oocyst polar granule present. Oocyst and sporocyst residua present. Sporocysts 8-10 x6 u. Form from M. tamariscinus: The following description is from Svan- baev (1962): Oocysts ellipsoidal or subspherical, 19-21 x 17-18 u, with a mean of 2117.5 u; length-width ratio 1.2. Oocyst wall smooth, yellow- orange or yellow-brown, 0.9-1.3 u thick. Micropyle absent. Oocyst polar granule usually present. Oocyst residuum a large, granular sphere be- tween the sporoblasts. Sporocysts ellipsoidal or subspherical, 9-13 x 6-8 u, with a mean of 11 x7 u. Sporozoites comma-shaped, 5—7 x 2-4 u, with a mean of 6x3 u. Sporocyst residuum in form of small granules. Schizogony and Gametogony: Presumably unknown. Prepatent Period: Presumably unknown. 88 THE COCCIDIAN PARASITES OF RODENTS Type Host: Meriones unguiculatus (Mongolian gerbil). Other Host: Meriones tamariscinus (tamarisk gerbil). Location: Presumably intestine. Geographic Distribution: USSR (Buryat Mongolia, Kazakhstan). Pathogenicity: Presumably unknown. Cross-Transmission Studies: Presumably none. Prevalence: Svanbaev (1962) found this species in one out of seven M. tamariscinus from southern Kazakhstan. Remarks: Machul’skii’s (1949) paper is not obtainable in this country, and the above description of the form from M. unguiculatus is taken from Svanbaev (1956) and Musaev and Veisov (1960). EIMERIA MARKOVI SVANBAEYV, 1956 (Blates35, Kig-3297) Eimeria markovi Svanbaev, 1956: 180-191. Description: Oocysts ovoid, 22-48 x 21-35 u, with a mean of 32.4 x 25.9 u. Oocyst length-width ratio 1.1—1.4, with a mean of 1.25. Oocyst wall smooth, greenish to yellowish green, with a double contour (but il- lustrated as a single layer), 1.4—-1.6 uw thick. Micropyle absent. Oocyst polar granule, oocyst residuum, and sporocyst residuum absent. Sporo- cysts described as ovoid but illustrated as ellipsoidal, 10-16 x 7-11 u, with a mean of 12.2 8.7 u. No Stieda body illustrated. Sporozoites comma- shaped, lying longitudinally in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones tamariscinus (tamarisk gerbil). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Trvansmission Studies: None. Prevalence: Svanbaev (1956) found this species in 23 per cent of 22 M. tamariscinus in western Kazakhstan. The same individuals were in- fected with FE. tamariscint. EIMERIA TAMARISCINI N. SP. Eimeria musculi Yakimoff and Gousseff. Svanbaey, 1956: 180-191 (from Meriones tamariscinus). [non] Eimeria musculi Yakimoff and Gousseff, 1938: 1-3; Svanbaev, 1956: 180-191 (from Apodemus sylvaticus and Microtus arvalis). Description: Oocysts subspherical, 19.1 x 18.6 uw in diameter. Length- width ratio 1.03. Oocyst wall smooth, greenish, double-contoured, 1.4 u thick. Micropyle absent. Oocyst polar granule absent. Oocyst and sporo- cyst residua absent. Sporocysts ovoid, 6.6 x 4.1 u. GENUS Eimeria SCHNEIDER, 1875 89 Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones tamariscinus (tamarisk gerbil). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in 23 per cent of 22 M. tamariscinus in western Kazakhstan. The same individuals were in- fected with E. markovt. Remarks: Svanbaev (1956) described coccidia from western Kazakhstan under the name Eimeria musculi not only from its type host, Mus musculus, but also from Apodemus sylvaticus, Microtus arvalis, and Meriones tamariscinus. The form he described from M. tamariscinus had smaller oocysts than E. musculi, although it resembled it in those other characters which he mentioned. However, in view of the failure of various investigators to transmit Ezmeria from one genus of rodent to another even within the same subfamily, and in view of the fact that Merzones and Mus belong to different rodent families, it is considered best to assign a new specific name to the form from M. tamariscinus. EIMERIA PESCHANKAE N. SP. Eimeria kriygsmanni [sic] Yakimoff and Gousseff. Svanbaev, 1962: 23-39. [non] Eimeria krijgsmanni Yakimoft and Gousseff, 1938: 1-3. Description: Oocysts ellipsoidal, subspherical, or spherical, 19-27 x 19-24 u, with a mean of 22x21 u; length-width ratio 1.1. Oocyst wall smooth, 1.3—2.1 u thick, yellow-green or yellow-brown. Micropyle absent. Oocyst polar granule not always seen. Oocyst residuum absent. Sporocysts ellipsoidal or spherical, 7-12 x 6-9 u, with a mean of 9x8 u. Sporocyst residuum absent. Sporozoites comma-shaped. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones tamariscinus (tamarisk gerbil). Location: Feces. Geographic Distribution: USSR (Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962) found this species in two out of seven M. tamariscinus in southern Kazakhstan. Remarks: Svanbaev (1962) described this species under the name “Eimeria kriygsmanni” (a misspelling for E. krijgsmanni). However, the 90 THE COCCIDIAN PARASITES OF RODENTS latter species was originally described from Mus musculus, and, for the reasons given under £. tamariscini above, we consider it a different species and have named it Eimeria peschankae; peschanka is the Russian word for gerbil. EIMERIA ASSAENSIS N. SP. Eimeria callosphermophili [sic] Henry. Svanbaev, 1962: 23-39 (in Meriones tamariscinus). [ron] Eimeria callospermophili Henry, 1932a: 279-290. Description: Oocysts subspherical or spherical, 25-30 x 23-26 u, with a mean of 28x25 u. Length-width ratio 1.1-1.2, with a mean of 1.1. Oocyst wall smooth, 1.2-1.4 u thick, greenish, yellow-green, or yellow- brown. Micropyle absent. Oocyst polar granule present. Oocyst residuum present in form of a large, granular ball. Sporocysts subspherical or ellipsoidal, 11-13 x 8-10 u, with a mean of 11.59 u. Sporocyst residuum absent. Sporozoites comma-shaped. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones tamariscinus (tamarisk gerbil). Location: Feces. Geographic Distribution: USSR (Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962) found this species in one out of seven M. tamariscinus in southern Kazakhstan. Remarks: Svanbaev (1962) described this species under the name “Eimeria callosphermophili” (a misspelling for E. callospermophilt). However, the latter species was originally described from Spermophilus lateralis, and, for the reasons given under £. tamariscini above, we con- sider it a different species and have named it Eimeria assaensis; the species name is derived from Assa, the region in southern Kazakhstan where Svanbaev found this coccidium. EIMERIA NORASCHENICA MUSAEV AND VEISOV, 1960 (Plate 18, Fig. 126) Eimeria noraschenica Musaev and Veisov, 1960a: 67-75. Description: Oocysts ovoid, rarely subspherical. Oocyst wall smooth, colorless to yellowish, composed of a single layer 1 u thick. Micropyle absent. Seventy-three sporulated oocysts from six animals measured 16—28 x 14-26 u, with a mean of 23 x 21 u; their length-width ratios ranged GENUS Eimeria SCHNEIDER, 1875 9] from 1.0-1.3, with a mean of 1.1. Oocyst residuum and polar granule absent. Sporocysts ovoid or spherical. Ovoid sporocysts 6-12 4-10 u, with a mean of 107 u. Spherical sporocysts 6-12 uw in diameter, with a mean of 8 u. Stieda body illustrated as absent. Sporocyst residuum composed of small granules located mainly between the sporozoites. Sporozoites lemon-shaped, illustrated without internal globules. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones persicus (Persian jird). Location: Large intestine contents. Geographic Distribution: USSR (Norashen and Ordubad regions of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA SALASUZICA MUSAEV AND VEISOV, 1960 (Plate 18, Fig. 131) Eimeria salasuzica Musaev and Veisov, 1960a: 67-75. Description: Oocysts ovoid, sometimes subspherical. Oocyst wall rough, granulated, with jagged surface, dark brown, composed of a single layer 1.5 uw thick. Micropyle absent. Twenty-one sporulated oocysts 22-26 x 20-24 u, with a mean of 24x 21 u; their length-width ratios ranged from 1.0-1.2, with a mean of 1.1. Oocyst residuum homogeneous, ovoid, or spherical, 6—7.5 u in diameter, with a mean of 7 u. Oocyst polar granule present. Sporocysts ovoid, 10-13 x 7-8 u, with a mean of 127 u. Stieda body prominent. Sporocyst residuum composed of small granules. Sporocyst refractile granule present. Sporozoite shape variable. Sporo- zoites illustrated without internal globules. Sporulation Time: Not given. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones persicus (Persian jird). Location: Large intestine contents. Geographic Distribution: USSR (Shakhbuz region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. 92 THE COCCIDIAN PARASITES OF RODENTS EIMERIA DISAENSIS MUSAEV AND VEISOV, 1960 (Plate 18, Figs. 127 and 128) Eimeria disaensis Musaev and Veisov, 1960a: 67-75. Description: Oocysts ovoid or subspherical. Oocyst wall smooth, color- less, composed of a single layer 1.0-1.2 uw thick. Micropyle prominent; Musaev and Veisov (1960a) found a small caplike body in the center of the micropyle of three oocysts. Fifteen sporulated oocysts from one host measured 18-24 x 16-22 u, with a mean of 23x18 u; their length-width ratios ranged from 1.0-1.2, with a mean of 1.1. Oocyst residuum and polar granule absent. Sporocysts ovoid, rarely spherical, 8-12 x 6-10 u, with a mean of 11x9 u. Stieda body absent. Sporocyst residuum com- posed of small granules mainly located between the _ sporozoites. Sporozoites ovoid, with a small globule in the broad end. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones persicus (Persian jird). Location: Large intestine contents. Geographic Distribution: USSR (Ordubad region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA LERIKAENSIS MUSAEV AND VEISOV, 1960 (Plate 18, Figs. 129 and 130) Eimeria lerikaensis Musaev and Veisov, 1960a: 67-75. Description: Oocysts ovoid or spherical. Oocyst wall smooth, composed of two layers, the inner being dark yellow and 1.0 u thick and the outer one colorless and 0.5-1.0 uw thick. Micropyle absent. Ninety-seven sporulated oocysts from six animals were measured. Seventy ovoid oocysts measured 14-22 x 12-20 u, with a mean of 19X16 yu; their length-width ratios ranged from 1.1-1.3, with a mean of 1.14. Twenty-seven spherical oocysts measured 14—22 yw in diameter, with a mean of 19 u. Oocyst residuum absent. Oocyst polar granule present. Sporocysts spherical or ovoid; ovoid sporocysts 6-10 x 4—8 u, with a mean of 9x7 uw; spherical sporocysts 4-8 in diameter, with a mean of 7.7 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites piriform, illustrated without internal globules. Sporulation Time: Three to four days at 25-30C in 2.5% potassium dichromate solution. GENUS Eimeria SCHNEIER, 1875 93 Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones persicus (Persian jird). Location: Large intestine contents. Geographic Distribution: USSR (Lerik, Yardymlin, Shakhbuz, and Norashen regions of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA SCHACHTACHTIANA MUSAEV AND VEISOV, 1960 : (Plate 19, Fig. 133) Eimeria schachtachtiana Musaev and Veisov, 1960b: 79-85. Description: Oocyst ovoid (illustrated as ellipsoidal). Oocyst wall smooth, colorless, composed of a single layer 1.0-1.25 u thick. Micropyle absent. Forty-four sporulated oocysts from four animals measured 16-30 x 14-24 u, with a mean of 24x 20 u; their length-width ratios ranged from 1.0—-1.4, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid (illustrated as ellipsoidal), 6-16 x 4-10 u, with a mean of 11 x8 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites, illustrated at the ends of the sporocysts, with a prominent globule at the broad end. Sporulation Time: Two to three days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones shawi (syn., M. tristramt) (Shaw’s jird). Location: Large intestine contents. Geographic Distribution: USSR (Dzhul’fin and Norashen regions of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA NEHRAMAENSIS MUSAEV AND VEISOV, 1960 (Plate 19, Fig. 134) Eimeria nehramaensis Musaev and Veisov, 1960b: 79-85. Description: Oocysts subspherical or spherical. Oocyst wall smooth, colorless, composed of a single layer 1 u thick. Micropyle absent. Seventy- three sporulated oocysts were measured: the subspherical ones were 14-26 x 12-24 u, with a mean of 2018 u, and length-width ratios of 1.0-1.2, with a mean of 1.05; the spherical oocysts were 14-22 u, with a 94 THE COCCIDIAN PARASITES OF RODENTS mean of 18 u. Oocyst residuum oval, illustrated as homogeneous, 4-8 x 3-6 u, with a mean of 6x5 u; or spherical, 2.5-6 u in diameter, with a mean of 5 u. Oocyst polar granule present. Sporocysts ovoid, 5-12 x 5-8 u, with a mean of 9x7 u. Stieda body present. Sporocyst residuum present, composed of a mass of small granules situated between the sporozoites. Sporozoites comma-shaped, with a prominent globule at the broad end. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones shawi (syn., M. tristrami) (Shaw’s jird). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA DZHAHRIANA MUSAEV AND VEISOV, 1960 (Plate 19, Fig. 135) Eimeria dzhahriana Musaev and Veisov, 1960b: 79-85. Description: Oocysts spherical or subspherical. Oocyst wall rough, dark brown, with a tuberculated surface, composed of a single layer 1.5 u thick. Micropyle absent. Twenty-one sporulated oocysts from one animal were measured: eleven subspherical ones measured 19-23 x 17-20 u, with a mean of 21x19 u, and length-width ratios of 1.1-1.2, with a mean of 1.1; ten spherical oocysts were 17.5-20 u in diameter, with a mean of 19 u. Oocyst residuum granular, spherical, 2-5 u in diameter, with a mean of 4 u. Oocyst polar granule present. Sporocysts ovoid, 6-10 x 4-6 u, with a mean of 9x 5 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites vary in shape, with a refractile globule. Sporulation Time: Three days at 25-30C in 2.5% potassium dichro- mate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones shawi (syn., M. tristrami) (Shaw’s jird). Location: Large intestine contents. Geographic Distribution: USSR (village of Dzhagri, Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. GENUS Eimeria SCHNEIDER, 1875 95 EIMERIA ARAXENA MUSAEV AND VEISOV, 1960 (Plate 19, Fig. 136) Eimeria araxena Musaev and Veisov, 1960b: 79-85. Description: Oocysts ovoid. Oocyst wall rough, dark brown, composed of a single layer 1.5—2 u thick. Micropyle absent. Forty-three sporulated oocysts from one animal measured 20-26 x 16-20 u, with a mean of 23x19 u; their length-width ratios ranged from 1.1—1.4, with a mean of 1.2. Oocyst residuum homogeneous, ovoid, rarely spherical: the ovoid residua measure 6-8 x 5-6 u, with a mean of 7 x6 u; the spherical residua are 5-6 u in diameter, with a mean of 6 u. Oocyst polar granule present. Sporocysts ovoid, rarely ellipsoidal, 9-13 x 5-8 u, with a mean of 11 x6 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites piriform, with a refractile globule in the broad end. Sporulation Time: Three and one-half to four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones shawi (syn., M. tristrami) (Shaw’s jird). Location: Large intestine contents. Geographic Distribution: USSR (Araksa region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA ASTRACHANBAZARICA MUSAEV AND VEISOV, 1960 (Plate 19, Fig. 137) Eimeria astrachanbazarica Musaev and Veisov, 1960b: 79-85. Description: Oocysts subspherical, ovoid, rarely spherical. Oocyst wall smooth, composed of two layers of which the outer is colorless and 1| u thick and the inner yellowish and 1 u thick. Micropyle absent. One hundred and twenty-four sporulated oocysts measured 15-30 x 14-26 u, with a mean of 22x19 u; their length-width ratios ranged from 1.0-1.4, with a mean of 1.1. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid, 6-14 4-10 u, with a mean of 9X7 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites comma-shaped, rarely piriform, with a refractile globule in the broad end. Sporulation Time: Three to 3.5 days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones shawi (syn., M. tristramz) (Shaw’s jird). 96 THE COCCIDIAN PARASITES OF RODENTS Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR and Astrakhan- bazar and Yardymlin regions of Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA VINOGRADOVI VEISOV, 1961 (Plate 20, 139) Eimeria vinogradovi Veisov, 1961: 25-35. Description: Oocysts ovoid or ellipsoidal. Oocyst wall smooth, colorless, composed of a single layer 1.0-1.5 u thick. Micropyle absent. Three hundred and twenty-three sporulated oocysts measured 16-34 x 14-30 u, with a mean of 22x19 u; their length-width ratios ranged from 1.1—1.6, with a mean of 1.3. Oocyst residuum absent. Oocyst polar granule very rarely seen. Sporocysts ovoid, ellipsoidal, and rarely spherical. The ovoid and ellipsoidal sporocysts measured 6-16 4-12 u, with a mean of 129 u; the spherical sporocysts were 6—12 u in diameter, with a mean of 9 u. Stieda body absent. Sporocyst residuum composed of small gran- ules, mainly between the sporozoites. Sporozoites comma-shaped, ovoid, or occasionally irregular, with a refractile globule in the broad end. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA ZULFIAENSIS VEISOV, 1961 (Plate 19, Fig. 138) Eimeria zulfiaensis Veisov, 1961: 25-35. Description: Oocysts ovoid. Oocyst wall smooth, colorless, composed of a single layer 1.0-1.25 u thick. Micropyle absent. One hundred and eighty-five sporulated oocysts measured 17-32 x 16-28 u, with a means of 25x20 u; their length-width ratios ranged from 1.0-1.4, with a mean of 1.2. Oocyst residuum homogeneous, ovoid, or spherical: the ovoid residua measured 6-12 4-10 u, with a mean of 8x7 u; the spherical GENUS Eimeria SCHNEIDER, 1875 97 residua were 4—12 in diameter, with a mean of 7 u. Oocyst polar granule very rarely seen. Sporocysts ovoid, 6-13 x 5-10 u, with a mean of 11 x 7 {ve Stieda body usually prominent. Sporocyst residuum composed of small granules between the sporozoites. Sporozoites comma-shaped, with a refractile globule in the broad end. Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Shakhbuz, Dzhul’fin, Norashen, and Nakhichevan regions of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA SADARAKTICA VEISOV, 1961 (Plate 20, Fig. 141) Eimeria sadaraktica Veisov, 1961: 25-35. Description: Oocysts ovoid. Oocyst wall smooth, colorless, composed of a single layer 1.4 u thick. Micropyle present. Sixty-two sporulated oocysts measured 18-24 x 16-22 u, with a mean of 23x19 u; their length-width ratios ranged from 1.1—1.4, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid, 6-12 4-8 u, with a mean of 107 u. Stieda body absent. Sporocyst residuum composed of a group of small granules in the center of the sporocyst. Sporozoites comma-shaped, with a refractile globule in the broad end. Sporulation Time: Three days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (vicinity of village of Sadarak, Norashen region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA JURSCHUAENSIS VEISOV, 1961 (Plate 20, Fig. 140) Eimeria jurschuaensis Veisov, 1961: 25-35. 98 THE COCCIDIAN PARASITES OF RODENTS Description: Oocysts ovoid. Oocyst wall rough, dark brown, composed of a single layer 2.3 u thick, which appears segmented and contains granules. Micropyle absent. Twenty-six sporulated oocysts measured | 30-36 x 24-30 u, with a mean of 34x27 u; their length-width ratios ranged from 1.2—1.4, with a mean of 1.3. Oocyst residuum homogeneous, subspherical, 10-12 u in diameter, with a mean of 11 uy. Oocyst polar granule absent. Sporocysts ovoid, 12-16 x 8-12 u, with a mean of 15x 10 u. Stieda body absent. Sporocyst residuum composed of a group of small granules in the center of the sporocyst. Sporozoites comma-shaped, with a refractile globule in the broad end. Sporulation Time: Three days at 25-30C in 2.59% potassium dichro- mate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Geographic Distribution: USSR (vicinity of village of Yurchi, Norashen region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA MUSAJEVI VEISOV, 1961 (Plate 21, Fig. 145) Eimeria musajevi Veisov, 1961: 25-35. Description: Oocysts ovoid, illustrated as ellipsoidal. Oocyst wall rough, tuberculated, dark brown, composed of a single layer 1.5-1.8 u thick dotted with small granules. Micropyle absent. Eighty-five sporulated oocysts measured 21-36 x 19-30 u, with a mean of 28 x 25 u; their length- width ratios ranged from 1.0-1.4, with a mean of 1.3. Oocyst residuum granular, ovoid or spherical; the ovoid residua measure 6-12 5-10 u, with a mean of 9x8 u; the spherical residua are 6—12 u in diameter, with a mean of 10 u. Oocyst polar granule absent. Sporocysts ovoid, 8-14 x 5-11 u, with a mean of 129 u. Stieda body absent. Sporocyst residuum com- posed of small scattered graules. Sporozoites piriform, with a refractile globule in the broad end. Sporulation Time: Four days at 25—30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR, Azerbaidzhan SSR). GENUS Eimeria SCHNEIDER, 1875 99 Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA TASAKENDICA VEISOV, 1961 (Plate 21, Fig. 144) Eimeria tasakendica Veisov, 1961: 25-35. Description: Oocysts ovoid, illustrated as ellipsoidal. Oocyst wall rough, colorless, composed of a single layer 1.2—1.4 u thick, with cross-striations. Micropyle absent. Sixty-six sporulated oocysts measured 19-26 x 17-23 u, with a mean of 23x21 u; their length-width ratios ranged from 1.1-1.3, with a mean of 1.1. Oocyst residuum homogeneous, ovoid or spherical: the ovoid residua measure 6-8 5-7 u, with a mean of 7x6 u; the spherical ones are 6-8 u in diameter, with a mean of 7 u. Oocyst polar granule present. Sporocysts ovoid, 8-13 x 5-9 u, with a mean of 127 u. Stieda body prominent. Sporocyst residuum composed of a group of small granules in the center of the sporocyst. Sporozoites comma-shaped, with a refractile globule in the broad end. Sporulation Time: Two days at 25—-35C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan region and Tazakend and other villages in the Norashen region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA BISTRATUM VEISOV, 1961 (Plate 20, Fig. 142) Eimeria bistratum Veisov, 1961: 25-35. Description: Oocysts ovoid or spherical. Oocyst wall smooth, composed of two layers, the outer one colorless to light yellow and | u thick, the inner one yellow-brown and | u thick. Micropyle absent. Two hundred and sixty-three ovoid oocysts measure 14-28 x 12-26 u, with a mean of 20x19 u; their length-width ratios ranged from 1.1—1.4, with a mean of 1.25. One hundred and twenty-five spherical oocysts ranged from 14-24 u in diameter, with a mean of 19 u. Oocyst residuum and polar granule absent. Sporocysts ovoid or spherical: the ovoid ones measured 100 THE COCCIDIAN PARASITES OF RODENTS 6-12 x 4-10 u, with a mean of 9X7 u; the spherical ones ranged from 4-11 u in diameter, with a mean of 10 u. Stieda body absent. Sporocyst residuum composed of small granules located between the sporozoites. Sporozoites varied in shape, without a refractile globule. Sporulation Time: Two or three days at 25-30C in 2.5°% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERA ARABIANA VEISOV, 1961 (Plate 20, Fig. 143) Eimeria arabiana Veisov, 1961: 25-35. Description: Oocysts ovoid. Oocyst wall smooth, composed of two layers, the outer one colorless and 0.7 u thick, the inner one brownish and 1.5 u thick. Micropyle absent. One hundred and forty-seven sporu- lated oocysts measured 16-32 x 12-26 u, with a mean of 25x20 u; their length-width ratios ranged from 1.1—-1.4, with a mean of 1.25. Oocyst residuum granular, spherical, 6-10 u in diameter, with a mean of 8 u. Oocyst polar granule absent. Sporocysts ovoid or piriform, 6-14 x 4-10 u, with a mean of 107 u. Stieda body absent. Sporocyst residuum com- posed of small granules. Sporozoites varied in shape, without a refractile globule. Sporulation Time: Three days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan region and vicinity of village of Arabengidzha in the Norashen region of Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. GENUS Eimeria SCHNEIDER, 1875 101 EIMERIA POLJANSKII VEISOV, 1961 Emend. (Plate 22, Fig. 154) Eimeria poljanski Veisov, 1961: 25-35. Description: Oocysts spherical or ovoid. Oocyst wall rough, colorless, composed of two layers: the outer one containing small granules and 0.6 u thick; the inner one containing small granules, corrugated, and 1.6 uw thick. Micropyle absent. Forty-seven sporulated oocysts were measured. The ovoid ones were 30—34 « 28—32 u, with a mean of 33 x 32 u; their length-width ratios ranged from 1.0-1.1, with a mean of 1.1. The spherical ones were 28-34 u in diameter, with a mean of 32 u. Oocyst residuum homogeneous, spherical, 5-8 yu in diameter, with a mean of 7 u. Oocyst polar granule absent. Sporocysts ovoid, 10-14 6-10 u, with a mean of 139 u. Stieda body prominent. Sporocyst residuum composed of small granules located between the sporozoites. Sporozoites comma- shaped, with a refractile globule at the broad end. Sporulation Time: Four days at 25—-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones vinogradovi (Vinogradov gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Nakhichevan ASSR, Azerbaidzhan SSR). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. EIMERIA ERYTHROURICA MUSAEV AND ALIEVA, 1961 (Plate 21, Fig. 146) Eimeria erythrourica Musaev and Alieva, 1961: 53-59. Description: Oocysts usually ovoid, sometimes spherical. Oocyst wall smooth, bright yellow to bright crimson, composed of a single layer 1.0-2.0 uw thick. Micropyle absent. Three hundred and eighty-three sporulated oocysts from 92 specimens measured 14-32 x 12-26 u, with a mean of 21x18 u; their length-width ratios ranged from 1.1—1.6, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid or spherical. Ovoid sporocysts 6-14 4-10 u, with a mean of 97 u; spherical sporocysts 6-10 u in diameter, with a mean of 8 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites pear- or bean-shaped, with a prominent spherical globule in the broad end. 102 THE COCCIDIAN PARASITES OF RODENTS Sporulation Time: Two to four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones libycus (syn., M. erythrourus) (red-tailed gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Alieva (1961) found this species in 92 speci- mens from 315 M. libycus from seven localities in Azerbaidzhan. Remarks: E. erythrourica resembles E. schachtachtiana from Meriones shawi, but differs from it in the fact that its sporozoites lie lengthwise in the sporocysts rather than at their ends. It also resembles E. vino- gradovi from M. vinogradovi, but differs from it in oocyst color, in having a polar granule, and in having a longer sporulation time. EIMERIA SCHAMCHORICA MUSAEV AND ALIEVA, 1961 (Plate 21, Fig. 147) Eimeria schamchorica Musaev and Alieva, 1961: 53-59. Description: Oocysts ovoid, rarely spherical. Oocyst wall smooth, colorless or sometimes with a yellow-crimson tinge, composed of a single layer 1.0-2.0 u thick. Micropyle absent. One hundred and thirty-three sporulated oocysts from 41 specimens measured 16-32 x 14-28 u, with a mean of 24x 20.5 u; their length-width ratios ranged from 1.0—1.4, with a mean of 1.2. Oocyst residuum homogeneous, 4-12 u in diameter, with a mean of 8 u; spherical sporocysts 6-10 u in diameter, with a mean of 9 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites pear- or bean-shaped, with a globule in the broad end. Sporulation Time: Three to four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones libycus (syn., M. erythrourus) (red-tailed gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Alieva (1961) found this species in 41 speci- mens from 315 M. libycus from Shamkhor and four other regions of Azerbaidzhan. GENUS Eimeria SCHNEIDER, 1875 103 EIMERIA ACHBURUNICA MUSAEV AND ALIEVA, 1961 (Plate 21, Fig. 148) Eimeria achburunica Musaev and Alieva, 1961: 53-59. Description: Oocysts ovoid. Oocyst wall smooth, colorless, composed of a single layer 1.0-1.5 uw thick. Micropyle prominent. Thirty-one sporulated oocysts from 11 specimens measured 14—24 x 12-21 u, with a mean of 20X17 u; their length-width ratios ranged from 1.1—1.4, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule rarely present. Sporocysts ovoid, rarely spherical. Ovoid sporocysts 6-10 x 4-8 u, with a mean of 9X7 u; spherical sporocysts 6-8 u in diameter, with a mean of 8 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites bean-shaped, with a globule in the broad end. Sporulation Time: Unknown. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones libycus (syn., M. erythrourus) (red-tailed gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Alieva (1961) found this species in 11 speci- mens from 315 M. libycus from the village of Agburun in the Martunin region of Azerbaidzhan. Remarks: E. achburunica resembles E. sadaraktica from Meriones vinogradovi, but differs from it in normally lacking a polar granule. EIMERIA SUMGAITICA MUSAEV AND ALIEVA, 1961 (Plate 21, Fig. 149) Eimeria sumgaitica Musaevy and Alieva, 1961: 53-59. Description: Oocysts ovoid, sometimes subspherical. Oocyst wall smooth, 1.5-2.0 u thick, composed of two layers, of which the outer is bright yellow and the inner is dark yellow. Micropyle absent. Thirty- six sporulated oocysts from eight specimens measured 16-26 by 14-24 u, with a mean of 22x19 u; their length-width ratios ranged from 1.1-1.4, with a mean of 1.14. Oocyst residuum absent. Oocyst polar granule present in some. Sporocysts ovoid, rarely spherical. Ovoid sporocysts 6-10 x 4-8 u, with a mean of 9X7 u; spherical sporocysts 6-8 u in diameter, with a mean of 8 u. Stieda body absent. Sporocyst residuum composed of small granules. Sporozoites pear-shaped, with a small globule. 104 THE COCCIDIAN PARASITES OF RODENTS Sporulation Time: Three to four days at 25-30C in 2.5% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones libycus (syn., M. erythrourus) (red-tailed gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Alieva (1961) found this species in eight speci- mens from 315 M. libycus from Sumgait and three other regions of Azerbaidzhan. Remarks: E. sumgaitica closely resembles E. astrachanbazarica from Meriones shawi, differing only in that the oocyst polar granule is not always present and in the fact that the outer layer of the oocyst wall is yellow rather than colorless; further study may show that it is a synonym. E. sumgaitica also resembles E. lerikaensis from M. persicus, but differs in having globules in the sporozoites. EIMERIA MARTUNICA MUSAEV AND ALIEVA, 1961 (Plate 21, Fig. 150) Eimeria martunica Musaev and Alieva, 1961: 53-59. Description: Oocysts ovoid. Oocyst wall smooth, 2 u thick, composed of two layers, of which the outer is bright yellow and the inner is dark yellow. Micropyle absent. Fifteen sporulated oocysts from four specimens measured 18-34 x 16-28 u, with a mean of 26x 22 u; their length-width ratios ranged from 1.1-1.3, with a mean of 1.14. Oocyst residuum spherical, granular, 4-12 u in diameter, with a mean of 9 u. Oocyst polar granule absent. Sporocysts ovoid, 8-14 x 6-10 u, with a mean of 118 u. Stieda body observed in some sporocysts. Sporocyst residuum composed of small granules. Sporozoites pear- or bean-shaped, illustrated without globules. Sporulation Time: Three to four days at 25-30C in 2.5% potassium dichromatic solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones libycus (syn., M. erythrourus) (red-tailed gerbil). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Alieva (1961) found this species in four GENUS Eimeria SCHNEIDER, 1875 105 specimens from 315 M. libycus from Martunin and two other regions of Azerbaidzhan. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily TACHYORYCTINAE EIMERIA TACHYORYCTIS VAN DEN BERGHE AND CHARDOME, 1956 (Plate 22, Fig. 157) Eimeria tachyoryctis van den Berghe and Chardome, 1956a: 67-69. Description: Oocysts described as ovoid but illustrated as ellipsoidal, averaging 2317 u. Oocyst wall quite thick (illustrated as composed of a single layer), colorless, apparently very resistant to pressure. Micro- pyle absent. Oocyst residuum nearly 6 uw in diameter. Oocyst polar granule not mentioned. Sporocysts 128 u, illustrated as ovoid and without Stieda body. Sporocyst residuum dark, quite large, polar. Sporozoites 8.5 x 3.5 u, reniform. Sporulation Time: Four to six days in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: The sporont divides into two round, granular masses 11 u in diameter. Each of these subdivides to form two sporocysts, leaving an oocyst residuum. Type Host: Tachyoryctes ruandae (fuku). Location: Probably intestine. Geographic Distribution: Africa (Belgian Congo). Pathogenicity: Unknown. Cross-Transmission Studies: Attempts to infect two half-grown white rats and two mice failed. Prevalence: This species was found in about 3 per cent of the T. ruandae at Tshibati, Lwiro (Kivu), Belgian Congo. Host Suborder MYOMORPHA Host Superfamily MUROIDEA Host Family MURIDAE Host Subfamily MURINAE EIMERIA MURIS GALLI-VALERIO, 1932 Eimeria muris Galli-Valerio, 1932: 129-142. [non] Eimeria muris: Pellérdy, 1954: 187-191 (see E. hungaryensis); Cerna, 1962: 1-13 (see E. hungaryensis). Description: Oocysts ovoid, with micropyle borne on a slight projec- 106 THE COCCIDIAN PARASITES OF RODENTS tion at narrow end. Oocysts 2115 u, with a spherical sporont 12 u in diameter. Sporocysts ovoid, 96 u. Oocyst residuum and polar granule not mentioned. Sporocyst residuum present. Sporozoites piriform. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus (syn., Mus) sylvaticus (field mouse). Location: Intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Pellérdy (1954) reported E. muris from Apodemus flavicollis in Hungary, but his description does not agree with Galli-Valerio’s, and we are assigning it a new name (p. 108). The form reported by Cerna (1962) as E. muris was apparently the same as that described by Pellérdy. EIMERIA NAYE GALLI-VALERIO, 1940 Eimeria naye Galli-Valerio, 1940: 387-392. Description: Oocysts cylindroid, with one end convex and the other flattened. Micropyle at flattened end. Oocysts 18-21 x 12-13.5 u. Sporont 10.5 w in diameter. Sporocysts spherical, 6 u in diameter. Oocyst residuum and polar granule not mentioned. Sporocyst residuum present. Sporo- zoites piriform. Sporulation Time: Four days on moist filter paper. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus (syn., Mus) sylvaticus (field mouse). Location: Intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Galli-Valerio (1940) found this species in two out of seven A. sylvaticus in Switzerland. EIMERIA APIONODES PELLERDY, 1954 (Plate 22, Fig. 151) Eimeria apionodes Pellérdy, 1954: 187-191. Description: Oocysts short, piriform, somewhat tapered at both ends, 17-23 x 13-18 u, with a mean of 2017 u. Oocyst wall relatively delicate (“zart’”), pale, and smooth. No micropyle illustrated or mentioned. Sporont round, finely granular. Oocyst residuum absent or composed at most of a few refractile granules. Polar granule absent. Sporocysts 128 u. Sporocyst residuum coarsely granular, so large (often 6 u in GENUS Eimeria SCHNEIDER, 1875 107 diameter) that it deforms the sporocyst, giving it an irregular appear- ance. Sporocyst Stieda body not mentioned or illustrated. Sporulation Time: Two to four days at room temperature in 2% potassium dichromate. Schizogony and Gametogony: Not described. Both the schizonts and gamonts are found in the epithelial cells of the small intestine mucosa. Prepatent Period: Eight to ten days. Type Host: Apodemus flavicollis (field mouse). Location: Small intestine. Geographic Distribution: Europe (Hungary). Pathogenicity: Unknown. Heavy mixed infections with this species and E. hungaryensis, E. rugosa, and E. apodemi caused catarrhal enteritis. Cross-Transmission Studies: Pellérdy (1954) was unable to transmit this species to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, and Cricetus cricetus. Prevalence: Pellérdy (1954) found E. apionodes and/or E. hungaryensis, E. rugosa, and E. apodemi in 23 per cent of 239 A. flavicollis in Hungary. EIMERIA APODEMI PELLERDY, 1954 (Plate 22, Figs. 152 and 153) Eimeria apodemi Pellérdy, 1954: 187-191; Cerna and Daniel, 1956: 19-23; Cerna, 1962: 1-13. [non] Eimeria apodemi: RySavy, 1957: 331-336 (see E. rysavy?). Description: Oocysts broadly ellipsoidal, often asymmetrical, 21-27 x 15-22 u, with a mean of 24x20 u (23-30 x 20-26 u, according to Cerna and Daniel, 1956). Oocyst wall brownish, composed of two layers. The outer brown layer is often detached during concentration in glycerol, revealing a lighter inner layer. Micropyle absent. Sporont round, not completely filling oocyst. Oocyst residuum absent. Polar granule absent. Sporocysts 127 uw according to Cerna and Daniel (1956). Sporocyst residuum present, finely granular. Stieda body not mentioned or il- lustrated. Sporulation Time: Three to five days at room temperature in 2% potassium dichromate. Schizogony and Gametogony: Not described. Prepatent Period: Six to seven days. Type Host: Apodemus flavicollis (field mouse). Other Host: Apodemus sylvaticus (field mouse). Location: Small intestine. Geographic Distribution: Europe (Hungary, Czechoslovakia). Pathogenicity: Unknown. Heavy mixed infections with this species and E. hungaryensis, E. rugosa, and E. apionodes caused catarrhal enteritis. 108 THE COCCIDIAN PARASITES OF RODENTS Cross-Transmission Studies: Pellérdy (1954) was unable to transmit this species to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, and Cricetus cricetus. Prevalence: Pellérdy (1954) found E. apodemi and/or E. hungaryensis, E. apionodes, and E. rugosa in 23 per cent of 239 A. flavicollis in Hungary. Remarks: RySavy (1957) reported finding E. apodemi in Clethrionomys glareolus in Czechoslovakia. Since, however, Pellérdy (1954) was unable to transmit E. apodemi from Apodemus to Clethrionomys, this identifica- tion cannot be correct, and we have named Ryéavy’s form E. rysavyi (see above). EIMERIA RUGOSA PELLERDY, 1954 (Plate 23, Fig. 158) Eimeria rugosa Pellérdy, 1954: 187-191. Description: Oocysts piriform, 23-27 x 15-19 u, with a mean of 24 x 16 u. Oocyst wall yellowish brown, somewhat thicker than usual. Under high magnification, the micropylar end appears wrinkled. Micropyle distinct. Sporont spherical, finely granular. Polar granule absent. Oocyst residuum large, usually more than 6 u in diameter. Sporocysts slender, 16 x 10 u. Sporocyst residuum small, finely granular. Sporulation Time: Two to four days at room temperature in 2% potassium dichromate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus flavicollis (field mouse). Location: Intestine. Geographic Distribution: Europe (Hungary). Pathogenicity: Unknown. Heavy mixed infections with this species and FE. hungaryensis, E. apionodes, and E. apodemi caused catarrhal enteritis. Cross-Transmission Studies: Pellérdy (1954) was unable to transmit this species to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, and Cricetus cricetus. Prevalence: Pellérdy (1954) found E. rugosa and/or E. hungaryensis, E. apionodes, and E. apodemi in 23 per cent of 239 4. flavicollis in Hungary. EIMERIA HUNGARYENSIS N. SP. (Plate 22, Figs. 155 and 156) Eimeria muris Galli-Valerio. Pellérdy, 1954: 187-191; Cerna, 1962: 1-13. [non] Eimeria muris Galli-Valerio, 1932: 129-142. Description: The following description was given by Pellérdy (1954) GENUS Eimeria SCHNEIDER, 1875 109 of the form in Apodemus flavicollis: Oocysts round or subspherical, 17-23 x 16-19 u, with a mean of 20x 18 u. Oocyst wall quite thick, light yellowish brown, with a somewhat rough surface. Sporont finely granular, completely filling unsporulated oocyst. Micropyle absent. Oocyst re- siduum and polar granule absent. Sporocysts 159 u, pointed at one end (apparently with a Stieda body) and rounded at the other. Sporocyst residuum present. The following description was given by Cerna (1962) for the form in A. sylvaticus: Oocysts broadly ellipsoidal (described as “oval’’) or spheri- cal, 14-24 x 14-16 u. Oocyst wall illustrated with a single layer. Micropyle absent. Oocyst polar granule and oocyst residuum absent. Sporocysts elongate ovoid, 9-13 4-8 u, with a Stieda body at the pointed end. Sporocyst residuum present. Sporulation Time: Two to three days in 2% potassium dichromate at room temperature according to Pellérdy (1954), four to five days according to Cerna (1962). Schizogony and Gametogony: Not described by Pellérdy (1954). Ac- cording to Cerna (1962), the sexual stages occurred in the epithelium of the posterior part of the small intestine (ileum). The mature micro- gametocytes measured 13-17 8-14 u, and the mature macrogametes measured 12-21 x 11-14 u. Prepatent Period: Four and one-half to five days. Type Host: Apodemus flavicollis (field mouse). Other Host: Apodemus sylvaticus (field mouse). Location: Mucosa of small intestine. Geographic Distribution: Europe (Hungary, Czechoslovakia). Pathogenicity: Unknown. Heavy mixed infections with this species and E. apionodes, E. apodemi, and E. rugosa were reported by Pellérdy (1954) to cause catarrhal enteritis. Cross-Transmission Studies: Pellérdy (1954) was unable to transmit this species to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, and Cricetus cricetus. Prevalence: Pellérdy (1954) found this species and/or E. apionodes, E. rugosa, and E. apodemi in 23 per cent of 239 A. flavicollis in Hungary. He remarked that it was the most common of the four species. Remarks: Pellérdy (1954) described this species under the name Eimeria muris Galli-Valerio, 1932, but it differs markedly from it in most of the few morphological features which Galli-Valerio mentioned. E. hungaryensis lacks a micropyle, it is less elongate than FE. murvs, its sporont completely fills the unsporulated oocyst, and its sporocysts are much larger. E. hungaryensis differs from E. naye Galli-Valerio, 1940 in lacking a micropyle, in being subspherical rather than cylindroid, and 110 THE COCCIDIAN PARASITES OF RODENTS in having elongate rather than spherical sporocysts. As shown by Pellérdy (1954), it also differs from E. apionodes, E. apodemi, and E. rugosa. We are therefore assigning it a new name. Cerna (1962) described what was probably the same species under the name FE. muris from Apodemus sylvaticus in Czechoslovakia. She also stated that the same form occurred in Mus musculus, but did not describe it; this identification is highly unlikely. EIMERIA PRASADLN. SP. Eimeria hindlei Yakimoft and Gousseff. Svanbaev, 1956: 180-191. [non] Eimeria hindlei Yakimoft and Gousseff, 1938: 1-3. Description: Oocysts ovoid, 26.2 x 19.8 u. Oocyst wall smooth, double- contoured, 1.6 u thick. Oocyst length-width ratio 1.32. Micropyle absent. Oocyst polar granule present (occasionally absent) in unsporulated oocyst. Oocyst residuum absent. Sporocysts ovoid, 8.86.6 u. Sporocyst residuum present. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus sylvaticus (common field mouse). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Svanbaev (1956) described this species under the name E. hindlei. He was unaware of Pellérdy’s (1954) descriptions of four species of Eimeria from A. flavicollis in Hungary or of Galli-Valerio’s (1932, 1940) descriptions of E. muris and E. naye, respectively, from A. sylvaticus in Switzerland. Pellérdy (1954) was unable to transmit any of his Apodemus coccidia to Mus musculus, Microtus arvalis, Clethrionomys (syn., Evotomys) glareolus, or Cricetus cricetus. Svanbaev’s description does not agree with those of the above six Eimeria species. His form differs from E. muris Galli-Valerio, 1932 in lacking a micropyle. It differs from E. naye Galli-Valerio, 1940 in shape, in lacking a micropyle, and in having ovoid rather than spherical sporocysts. It differs from E. apionodes Pellérdy, 1954 in shape; in addi- tion, it is larger but has smaller sporocysts. It differs from E. apodemi Pellérdy, 1954 in shape and color, and in having a polar granule. It differs from E. hungaryensis n. sp. (see above) in shape, in having a smooth, greenish wall rather than a thick, yellowish brown rough wall, in having a polar granule, and in having much smaller sporocysts. It differs from E. rugosa Pellérdy, 1954 in shape, in not having a yellowish brown wall wrinkled at one end, in having a polar granule, in lacking GENUS Eimeria SCHNEIDER, 1875 111 an oocyst residuum and a micropyle, and in having much smaller sporo- cysts. In view of these morphological differences and of the failure of Pellérdy’s cross-transmission attempts, it is considered best to assign this form a new name. EIMERIA SVANBAEVI N. SP. (Plate 23, Fig. 159) Eimeria kriygsmanni [sic] Yakimoff and Gousseff. Svanbaev, 1956: 180-191 (from A podemus sylvaticus). (?) Eimeria krijgsmant [sic]: RySavy, 1954: 131-174. [non] Eimeria krijgsmanni Yakimoft and Gousseff, 1938: 1-3. [non] Eimeria kriygsmanni [sic]: Svanbaev, 1956: 180-191 (from Mus musculus). Description: Oocysts ovoid, colorless or occasionally greenish, with a smooth, double-contoured wall 1.6—1.8 u thick, 24-26 x 20-22 u, with a mean of 24.9 20.5 u and a mean length-width ratio of 1.22. Micropyle absent. Oocyst polar granule present. Oocyst and sporocyst residua absent. Sporocysts ovoid, 9-13 x 7-9 u, with a mean of 10.3 x 7.7 u. Sporo- zoites comma-shaped. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus sylvaticus (field mouse). Location: Feces. Geographic Distribution USSR (western Kazakhstan), Czechoslovakia (?). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Svanbaev (1956) described this species under the name E. kriygsmanni Yakimoff and Gousseff, 1938 (lapsus for E. krijgsmanni). See Remarks under E. prasadi (p. 110) for a discussion of this paper. Svanbaev’s description does not agree with those of any other Apodemus coccidia. It differs from E. muris Galli-Valerio, 1932 in lack- ing a sporocyst residuum and a micropyle. It differs from E. naye Galli- Valerio, 1940 in shape and size, in lacking a micropyle and sporocyst residuum, and in shape of the sporocysts. It differs from E. apionodes Pellérdy, 1954 in shape, in having a thicker wall, and in lacking a sporocyst residuum. It differs from E. apodemi Pellérdy, 1954 in shape, in the color of the oocyst wall, in having an oocyst polar granule, and in lacking a sporocyst residuum. It differs from E. rugosa Pellérdy, 1954 in shape, in the character of the oocyst wall, in having an oocyst polar granule, and in lacking a micropyle and oocyst and sporocyst residua. It differs from E. hungaryensis n. sp. in shape, in the character of the oocyst wall, in having an oocyst polar granule, and in lacking a sporocyst 112 THE COCCIDIAN PARASITES OF RODENTS residuum. It differs from E. prasadi n. sp. in lacking a sporocyst residuum. In view of these morphological differences and of the failure of Pellérdy’s (1954) cross-transmission attempts, it is considered best to assign this form the name, Eimeria svanbaevi n. sp. RySavy (1954) described an Eimeria from the intestine of A. sylvaticus in Czechoslovakia under the name FE. krijgsmani (lapsus for E. krijgs- manni) Yakimoff and Gousseff, 1938. He, too, was unaware of the papers of Galli-Valerio and Pellérdy cited above. The oocysts of his form were described as oval, broadly rounded at either end, and stretched out; they were illustrated as ellipsoidal. They measured 16-27 x 12-16 u, with a mean of 20.2x15.1 u. Their length-width ratios ranged from 1.0-1.8, with a mean of 1.35. The oocyst wall was thin, colorless, and easily broken. There was no micropyle. The sporocysts measured 10-12 x 5-6 u. No information was given on the oocyst or sporocyst residua or the polar granule. This form appears to resemble E. svanbaevi more than the other species of Ez:meria described from Apodemus, although it differs from it in some respects. However, until a better description is available, we do not feel justified in establishing a new name for it, and for the present assign it with a question mark to EF. svanbaevi. EIMERIA RUSSIENSIS N. SP. Eimeria musculi Yakimoff and Gousseff. Svanbaev, 1956: 180-191 (from A podemus sylvaticus). [non] Eimeria musculi Yakimoff and Gousseff, 1938: 1-3; Svanbaev, 1956: 180-191 (from Mus musculus—see E. musculi) (from Microtus arvalis—see E. arvicolae) (from Meriones tamariscinus—see E. tamariscint). Description: Oocysts spherical, greenish, smooth, with a double-con- toured wall 1.5—-2.0 u thick, 21.7 » in diameter. Micropyle absent. Polar granule present, occasionally absent. Oocyst and sporocyst residua absent. Sporocysts oval or spherical, 8.78.6 u. Sporozoites comma-shaped, 6x4 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus sylvaticus (field mouse). Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Svanbaev (1956) described this species under the name E. muscult Yakimoff and Gousseff, 1938. See Remarks under E. prasadi (p. 110) for a discussion of his paper. Svanbaev’s description does not agree with those of any of the Apodemus coccidia so far named. It differs from E. murts Galli-Valerio, 1932 and from E. naye Galli-Valerio, 1940 GENUS Eimeria SCHNEIDER, 1875 113 in shape and in lacking a sporocyst residuum and a micropyle. It differs from E. apionodes Pellérdy, 1954 in oocyst and sporocyst shape and in lacking a sporocyst residuum. It differs from FE. apodemi Pellérdy, 1954 in oocyst shape. in the character of the oocyst wall, in lacking a sporocyst residuum, and in having a polar granule. It differs from E. rugosa Pellérdy, 1954 in oocyst shape, in the character of the oocyst wall, in having a polar granule, and in lacking a micropyle and oocyst and sporocyst residua. It differs from FE. hungaryensis n. sp. in the character of the oocyst wall, in having a polar granule, in lacking a sporocyst residuum, and in the shape of the sporocysts. It differs from E. prasadi n. sp. in oocyst and sporocyst shape and in lacking a sporocyst residuum. It differs from FE. svanbaevi n. sp. in oocyst and sporocyst shape. In view of these morphological differences and of the failure of Pellérdy’s (1954) cross-transmission attempts, it is considered best to assign this form the name Eimeria russiensis n. sp. EIMERIA SP. (RYSAVY, 1954) Eimeria falciformis (Eimer). Elton et al., 1931: 657-721; RySavy, 1954: 131-174 (pro parte); Cerna and Daniel, 1956: 19-23 (pro parte). [non] Eimeria falciformis (Eimer, 1870) Schneider, 1875: xl-xlv. Description: Uncertain. Hosts: Apodemus sylvaticus, A. flavicollis (field mice). Location: Feces. Geographic Distribution: England, Czechoslovakia. Remarks: RySavy (1954) reported E. falciformis from Mus musculus, Apodemus flavicollis, A. sylvaticus, Microtus arvalis, and Clethrionomys glareolus in Czechoslovakia. The description he gave was similar to that of E. falciformis. For the reasons given in the discussion of these forms in Microtus arvalis (p. 81), it is very doubtful that the forms RySsavy saw in the hosts other than M. musculus were E. falciformis. However, since he did not describe them separately from each host, it is useless to attempt to assign names to them. Cerna and Daniel (1956) reported EF. falciformis from Apodemus flavicollis and Clethrionomys glareolus in Czechoslovakia. They gave the same dimensions as RySavy, but mentioned that the oocyst wall was more than 1 u thick. For the reasons given above, these forms were undoubtedly not EF. falciformis, but cannot be assigned separate names. Elton et al. (1931) reported finding E. falciformis in 38 per cent of 380 wild Apodemus sylvaticus in England. They did not describe the oocysts. These coccidia were undoubtedly not E. falciformis, which is a parasite of the house mouse and not of Apodemus, but cannot be as- signed to any species. 114 THE COCCIDIAN PARASITES OF RODENTS EIMERIA SP. (RYSAVY, 1954) (Plate 23, Figs. 160 and 161) Eimeria keilini Yakimoft and Gousseff. RySavy, 1954: 131-174; Cerna and Daniel, 1956: 19-23; Cerna, 1962: 1-13. [non] Eimeria keilint Yakimoff and Gousseff, 1938: 1-3. Description: Oocysts described by RySavy (1954) from Apodemus sylvaticus as ellipsoidal, narrow at both ends. Oocyst wall thin, trans- parent, colorless, composed of a single layer. Oocysts 24-29 x 16-20 u, with a mean of 26.0 x 20.7 [sic] u. Micropyle absent. The oocysts did not sporulate. Cerna and Daniel (1956) described what was probably the same form from 4A. flavicollis in Czechoslovakia. Its oocysts measured 25-28 x 16-20 u, were narrow at both ends, and had a thin, colorless wall and no micropyle. The sporocysts measured 8-9 x 6-7 u. Nothing was said about oocyst or sporocyst residua or oocyst polar granules, but these were not visible in the rather hazy photomicrograph. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus sylvaticus (field mouse). Other Host: Apodemus flavicollis (field mouse). Location: Intestine. Geographic Distribution: Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This form was found by RySsavy (1954) in 2 A. sylvaticus and by Gerna and Daniel (1956) in 1 out of 16 A. flavicollis. Remarks: Rysavy (1954) described this form under the name E. kezlini. See Remarks under Eimeria sp. from Microtus arvalis (p. 81) for a discussion of his paper. RySavy’s oocysts did not sporulate. Cerna and Daniel (1956) and Cerna (1962) found the same form in Apodemus flavicollis in Czechoslovakia. Their oocysts sporulated. The sporocysts were smaller than those of E. keilini, which measure 12 x 6 u. In addition, its wall was colorless rather than yellowish. In view of these differences and of the failure of Pellérdy’s (1954) cross-transmission experiments, it is unlikely that RySavy’s form is EF. keilini, whose type host is Mus musculus. Its shape is unlike that of any of the species of Eimeria so far described from Apodemus, but in the absence of more information it is not considered desirable to assign it a specific name. EIMERIA (2) SP. (RYSAVY, 1954) Eimeria hindlei Yakimoft and Gousseff. RySavy, 1954: 131-174 (pro parte). [non] Eimeria hindlei Yakimoff and Gousseff, 1938: 1-3. Description: Oocysts oval. Oocyst wall thin, pale yellow, composed GENUS Eimeria SCHNEIDER, 1875 115 of a single layer. Micropyle absent. Oocysts 23-27 x 18-20 u, with a mean of 25.0 18.4 u. Oocyst length-width ratio 1.2-1.5, most often 1.4. The oocysts did not sporulate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus sylvaticus (field mouse). Location: Feces. Geographic Distribution: Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: RySavy (1954) described this form under the name E. hindlei from A. sylvaticus and Clethrionomys glareolus in Czechoslovakia. He did not differentiate between the forms from the two hosts and the oocysts failed to sporulate. See Remarks under Eimeria sp. from Microtus arvalis (p. 81) for a discussion of his paper. Since the type host of EF. hindlei is Mus musculus, it is clear that RySavy’s form does not belong to this species. As a matter of fact, it is unlikely that RySavy’s forms from 4. sylvaticus and C. glareolus belong to the same species. Furthermore, since they did not sporulate, they may not even be Eimeria. EIMERIA (?) SP. CERNA AND DANIEL, 1956 Eimeria sp. Cerna and Daniel, 1956: 19-23. Description: Oocysts broadly ovoid to spherical, 11-14 x 10-13 u. Oocyst wall very delicate. Micropyle absent. The oocysts failed to sporulate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Apodemus flavicollis and Clethrionomys glareolus. Location: Intestine. Geographic Distribution: Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Cerna and Daniel (1956) found this form in 1 out of 16 A. flavicollis and 1 out of 31 C. glareolus. Remarks: Although Cerna and Daniel (1956) assigned this form to Eimeria, its failure to sporulate makes it impossible to be certain of its genus. EIMERIA VINCKEI RODHAIN, 1954 (Plate 37, Fig. 300) Eimeria vinchei Rodhain, 1954: 327-329. Description: Oocysts cylindroid, colorless, with a thin, double-con- toured wall, 20-24 x 12-15 u, with a mean of 22.0 x 13.2 u. Length-width 116 THE COCCIDIAN PARASITES OF RODENTS ratio 1.7. Micropyle absent. Sporont in unsporulated oocysts 13.2 u in diameter. Oocyst residuum absent. Oocyst polar granule not mentioned or illustrated. Sporocysts 7x4 u (illustrated as round), apparently with- out Stieda body. Sporocyst residuum present. Sporulation Time: Seven to eight days. Schizogony and Gametogony: Six to ten merozoites are produced by schizogony. The mature macrogametes are round and about 9 u in diam- eter. The microgametocytes are oval, 149 u. Prepatent Period: Eleven days. Type Host: Thamnomys surdaster surdaster. Location: Cecum. The parasites are found in the superficial epithelial cells bordering the folds of the mucosa. Geographic Distribution: Africa (Haut Katanga, Belgian Congo). Pathogenicity: Nonpathogenic. Cross-Transmission Studies: Rodhain (1954) was unable to transmit this species to laboratory mice or rats. Prevalence: Rodhain (1954) found E. vinckei in 12 out of 38 T. s. surdaster in Haut Katanga. EIMERIA DASYMYSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 * (Plate 23, Fig. 162) Eimeria dasymysis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts subspherical, ellipsoidal, sometimes slightly ovoid. Oocyst wall colorless to pale yellowish, smooth, composed of a single layer about 0.8 u thick. Micropyle absent. Thirty-eight oocysts from one host animal measured 17-23 x 15-21 u, with a mean of 19.8x 17.0 u; their length-width ratios ranged from 1.0-1.4, with a mean of 1.16. Oocyst polar granule present. Oocyst residuum absent. Sporocysts 10-11 x 6 u; sporocyst length-width ratio 1.6—-1.9, with a mean of 1.75. Sporocysts slightly ovoid, with a tiny, buttonlike Stieda body. Sporocyst wall very thin. Sporocyst residuum present, granular. Sporozoites homogeneous, more or less curled up in sporocysts. Sporulation Time: Three days at room temperature in Liberia. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dasymys incomptus rufulus. Location: Lower jejunum. The macrogametes and immature parasites lie in the epithelial cells below the host cell nuclei (i.e., distal to the lumen). * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 nnl7, Geographic Distribution: Liberia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine, Bray, Ivens, and Gunders (1959) found this species in two animals in Harbel, Liberia. EIMERIA ARVICANTHIS VAN DEN BERGHE AND CHARDOME, 1956 (Plate 37, Fig. 301) Eimeria arvicanthis van den Berghe and Chardome, 1956: 65-66. Description: Oocysts described as ovoid but illustrated as ellipsoidal, 23-24 x 10-14 u. Oocyst wall thin, rose-colored, number of layers not given. Micropyle presumably absent. Oocyst residuum composed of some rare granules. Oocyst polar body not mentioned, presumably absent. Sporocysts 10-117 u, illustrated as ellipsoidal without Stieda body. Sporocyst residuum composed of about 20 granules arranged in a rosette. Sporozoites 7 x 3 u, with one end broader than the other. Sporulation Time: Seventy-eight hours in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicanthis abyssinicus rubescens. Location: Probably intestine. Geographic Distribution: Africa (Belgian Congo). The animal(s?) was captured at Tshibati (Lwiro), Kivu. Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA PUTEVELATA BRAY, 1958 (Plate 23, Fig. 164) Eimeria putevelata Bray, 1958: 81-83. Description: Oocysts ovoid, 22-30 x 17-22 u, with a mean of 26.5x 19.9 u; mean length-width ratio, 1.26. Oocyst wall composed of a thick, yellow outer layer covered with small, well-defined pits and a thin inner layer. Micropyle absent. Occasional oocyst polar granules present. Oocyst residuum absent. Sporocysts 10-13 x 8-10 u, with a mean of 11.5 8.3 u, ovoid, with a very small Stieda body which is only a slight thickening of the wall at the narrow end. Sporocyst residuum usually but not always present. Sporozoites 10x 3 u, banana-shaped. Sporulation Time: Ten days. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lemniscomys striatus striatus (striped grass mouse). 118 THE COCCIDIAN PARASITES OF RODENTS Location: Yleum. Bray (1958) found macrogametes in the epithelial cells of the posterior third of the small intestine. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA LEMNISCOMYSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 * (Plate 23, Fig. 163) Eimeria lemniscomysis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts broadly spindle-shaped with somewhat flattened ends. Oocyst wall brownish yellow, moderately rough and pitted, com- posed of a single layer about 1.2 u thick at the sides and 0.8 u thick at the ends. A thin membrane lines the wall. Micropyle absent. Three sporulated oocysts measured 27-30 x 18-19 u, with a mean of 28.3 x 18.8 u; their length-width ratios were 1.5-1.6. Oocyst polar granule present. Oocyst residuum absent. Sporocysts 168 u, elongate ovoid, pointed at one end; Stieda body small or absent. Sporocyst residuum present, finely granular. Sporozoites lie more or less diagonally in sporocyst. Oocysts collapse rather quickly in Sheather’s sugar solution. Sporulation Time: Three days at room temperature in Liberia. Schizogony and Gametogony: Macrogametes contain eosinophilic gran- ules concentrated at their periphery. Prepatent Period: Unknown. Type Host: Lemniscomys striatus striatus (striped grass mouse). Location: Macrogametes and microgametocytes in epithelial cells of upper jejunum and lower duodenum, lying between host cell nucleus and lumen. Geographic Distribution: Liberia. Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA MIYAIRII OHIRA, 1912 (Plate 23, Figs. 165 and 166; Plate 24, Figs. 167-181; Plate 37, Figs. 302-312) Eimeria miyairit Ohira, 1912: 1045-55; Roudabush, 1937: 135-163. Eimeria myiari [sic]: RySavy, 1954: 131-174. Eimeria carinii Pinto, 1928: 127-128; Becker and Burroughs, 1933: 123; Becker, 1934; Matubayasi, 1938: 144-163. [non] Eimeria miyairi: Pérard, 1926: 120-124; Becker, Hall, and Hager, 1932: 299-316; Becker, 1934; Yakimoff and Gousseff, 1935: 261-265. Description: Oocysts spherical to subspherical, with a thick, rough, * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 119 yellowish brown, radially striated wall. According to Matubayasi (1938), the wall is composed of two layers, the outer one thin and rough, and the inner one thick and radially striated. On the basis of Pinto’s (1928) illustration, there may be two layers in the oocyst wall. Oocysts 17-29 x 16—26 u, with a mean of 24x22 u and a mean length-width ratio of 1.10, according to Becker and Burroughs (1933). Micropyle absent. Oocyst polar granule and oocyst residuum absent. Sporocysts 16-18 9-10 u according to Pinto (1928). Sporocyst residuum present. Sporulation Time: Four to five days (six days, according to Matubayasi, 1938). Schizogony: Roudabush (1937) described the endogenous stages of E. miyairii in detail. The sporozoites are most abundant in the small in- testine about 12 hours after infection. They are banana-shaped, 12-17 u long (mean, 14.5 u) and 2-3 u wide, with a siderophilic, refractile globule at each end. The nucleus has a marginal chromatic ring and a central karyosome, and lies anterior to the middle of the sporozoite. The sporozoite enters an intestinal cell and rounds up to form a first- generation schizont. This produces 12 to 24 first-generation merozoites and a residual mass. ‘These merozoites measure 6—7 x 1—2 u, with a mean of 6.6 x 1.3 u. They have a central nucleus with a central karyosome, but no refractile globules. They leave the host cell in two days, enter new host cells, and form second-generation schizonts. Each second-generation schizont produces 8 to 16 second-generation merozoites plus a residuum on the third day. ‘These merozoites measure 8-11 x 1-2 u, with a mean of 9.2 1.4 u. The nucleus is in the posterior fourth of the body; there are usually one or two large granules anterior to it. The second-generation merozoites enter new host cells and form third-generation schizonts. These produce 20 to 24 third-generation merozoites and a large re- siduum on the fourth day of infection. These merozoites measure 4—5 x 1-2 u, with a mean of 4.4 1.2 u, and have a central nucleus. Gametogony: The third-generation merozoites enter new host cells. Roudabush (1937) considered that they formed microgametocytes and macrogametocytes, but the cells he considered macrogametocytes are actually macrogametes. The young “gametocytes” are indistinguishable and have a central vacuolated area containing an eccentric chromatin mass. The microgametocyte nucleus divides repeatedly, and the nuclei thereby produced migrate to the periphery of the cell, where the micro- gametes form. A large residual body is left. The microgametes measure 30.75 uw and have two flagella at their anterior end. As the macrogametes develop, large, eosinophilic plastic granules and very small hematoxylinophilic granules form in their cytoplasm. The plastic granules migrate to the periphery, and fuse to form the outer wall of the oocyst. Roudabush (1937) found microgametes in the tissues 120 THE COCCIDIAN PARASITES OF RODENTS around these developing macrogametes, and believed that fertilization takes place while the latter are still in their host cells. Mature oocysts appear 5.5 days after infection. Roudabush (1937) calculated that E. miyairii could produce 25,360 to 73,728 (mean, 38,016) oocysts per oocyst fed. According to Becker and Burroughs (1933), oocysts continue to be discharged for five days after a single inoculation, and for six to seven days after repeated inoculations. The average number of oocysts discharged during the patent period following repeated inoculation was 1.99 x 10°. Prepatent Period: Six days (six to eight days, according to Matubayasi, 1938). Type Host: Rattus norvegicus (Norway rat). Other Host: Rattus rattus (2). Matubayasi (1938) found this species in 4 of 33 wild rats (R. norvegicus and R. rattus) which he examined in Japan. However, he did not specify whether it occurred in one or both host species. Location: Small intestine. E. miyairii occurs almost exclusively in the epithelial cells of the villi, but has been found occasionally in the glands. A striking difference between it and other rat coccidia is its tendency to cause parasitized cells to be pushed out of the epithelial layer toward the tunica propria. In addition, it frequently causes double or even multiple infections of cells, especially in the later stages of its life cycle. Roudabush’s (1937) figures show that the parasites may be found beneath, above, or beside the host cell nucleus. Ohira’s (1912) figures indicate that they are usually beneath the host cell nucleus. Geographic Distribution: Worldwide. Pathogenicity: Apparently unknown. Cross-Transmission Studies: None. Remarks: The synonymy of this species indicates the confusion which has existed regarding its correct name. The first authentic description of a coccidium from the rat was Ohira’s (1912) description of E. miyairit. Ohira described only the endogenous stages and did not see the oocysts. His description was in Japanese, and was not available to most workers, who were guided by a German abstract of his paper. Pérard (1926) accepted the name EF. miyairit, for the coccidium with which he worked, believing that EF. nieschulzi Dieben, 1924 was a synonym. Subsequent workers followed his lead. Pinto (1928) described the oocysts of E. miyairi for the first time; he believed it to be a new species to which he gave the name, E. carinii. Roudabush (1937) finally straightened out the nomenclatural tangle. He studied the endogenous stages of the three commonest rat coccidia, compared them with the figures in Ohira’s (1912) paper, and concluded that Pérard’s E. miyairii was actually FE. GENUS Eimeria SCHNEIDER, 1875 121 nieschulzi, while Pinto’s E. carinii was actually E. miyairii. We ourselves obtained a copy of Ohira’s (1912) paper, had it translated, and are able to confirm Roudabush’s interpretation. Without being aware of Rouda- bush’s paper, Matubayasi (1938) made a study of this species and came to the same conclusion, but nevertheless continued to use the name E. carinii for E. miyairit and the name of E. miyairi for E. nieschulzi. EIMERIA NIESCHULZI] DIEBEN, 1924 (Plate 25, Figs. 182-187; Plate 26, Figs. 194-208; Plate 27, Figs. 209-214; Plate 38, Figs. 313-325; Plate 47, Figs. 438-443; Plate 48, Figs. 444-449) Eimeria nieschulz Dieben, 1924; Roudabush, 1937: 135-163. Eimeria falciformis (Eimer) Schneider pro parte. Auctores. Eimeria miyairii Ohira. Pérard, 1926: 120-124; Becker, Hall, and Hager, 1932: 299-316; Becker, 1934; Yakimoff and Gousseff, 1935: 261-265; Matubayasi, 1938: 144-163; Beltran and Pérez, 1950: 71-78. Eimeria mytairii [sic]: Yakimoff and Gousseff, 1936a: 504-508. Eimeria halli Yakimoff, 1935: 81=83; Yakimoff and Gousseff, 1936a: 504-508. Description: Oocysts ellipsoidal to ovoid, tapering at both ends. Oocyst wall smooth or granular, colorless to yellowish, composed of a single layer about 1.1 u thick. Becker, Hall, and Hager (1932) illustrated the wall as composed of two layers, and Becker (1934) and Dieben (1924) stated that it was composed of two layers; on the other hand, Matubayasi (1938), reported only one layer, Pérard (1926) illustrated a single layer, and we ourselves found only a single layer in the forms from R. norvegicus and R. hawaiiensis which we studied. The oocysts described by Becker, Hall, and Hager (1932) from Iowa rats measured 16-26 x 13-21 u, with a mean of 22.5 17.8 u; their length-width ratios ranged from 1.0—1.5, with a mean of 1.26. Sixteen oocysts from a wild R. norvegicus in Illinois measured 18-24 15-17 u, with a mean of 20.7 16.5 u; their length- width ratios ranged from 1.1—1.4, with a mean of 1.25. Oocyst polar granule present. Oocyst residuum absent. Sporocysts elongate ovoid, with very small Stieda body. Sporocyst wall thin. Five sporocysts from a wild R. norvegicus in Illinois measured 11-12 x 7 u, with a mean of 11.8 x 7.0 u; their length-width ratios ranged from 1.6—1.8, with a mean of 1.64. Sporo- cyst residuum present, compact. Sporozoites lie lengthwise, head to tail, in sporocysts. Sporozoites with a pale yellowish, clear globule at the large end. Sporulation Time: Two and one-half to three days (two days accord- ing to Matubayasi, 1938; three and one-half days according to Dieben, hg? 4). Schizogony: Roudabush (1937) described the endogenous stages of E. nieschulzi in detail. The sporozoites may be found in the lumen of the small intestine three to four hours after infection, and may remain in 122 THE COCCIDIAN PARASITES OF RODENTS the intestine in the infective condition for as long as four days. They measure 10-12 x 1-2 u, with a mean of 11.3 x 1.8 u, and have a siderophilic, eosinophilic refractile globule at either end. The nucleus has a central karyosome. The sporozoite enters an intestinal cell and rounds up to form a first- generation schizont; this schizont differs from those of later generations in containing a large, refractile, eosinophilic globule. The first-genera- tion schizont produces 20 to 36 (mean, 26) first-generation merozoites in about 36 hours. These merozoites measure 7-10 x 1-2 u, with a mean of 8.6 x 1.6 u, and have a small eosinophilic (but not siderophilic) globule at either end. They break out of the host cell and enter other intestinal cells, where they round up to form second-generation schizonts. These mature about 48 hours after infection, forming 10 to 14 (mean, 12) second-generation merozoites which measure 13-16 1 u, with a mean of 14.4x1.2 u; these do not have eosinophilic globules. The second- generation merozoites break out of the host cell and enter new intestinal cells, where they round up to form third-generation schizonts which mature about 72 hours after infection, giving rise to 8 to 20 (mean, 15) third-generation merozoites. These measure 17-22 x1 u, with a mean of 19.11.2 u, and are usually U- or J-shaped. The third-generation merozoites enter new intestinal cells, where they round up to form fourth-generation schizonts. These mature in four days after infection, giving rise to 36 to 60 (mean, 50) fourth-generation merozoites which measure 4—7 x 1-2 u, with a mean of 5.5x 1.4 u. Gametogony: The fourth-generation merozoites enter new intestinal cells. Roudabush (1937) considered that they formed microgametocytes and macrogametocytes, but the cells he called macrogametocytes are actually macrogametes. The young “gametocytes” are first seen about 5.5 days after infection, but cannot be distinguished until later. They consist of a central mass of chromatin surrounded by a crescentic, non- granular area which is in turn surrounded by rather densely granular cytoplasm. The microgametocyte nucleus divides repeatedly, the nuclei migrate to the periphery of the gametocyte, their chromatin becomes elongate, and they form comma-shaped microgametes with two anterior flagella. ‘The microgametes measure 4.40.1 u, and their flagella are 9.2 u long. The macrogametes contain two types of granules. The larger ones are eosinophilic and the smaller ones are basophilic and hematoxy- linophilic. The eosinophilic granules, known as plastic granules, migrate to the periphery of the gamete, where they flatten out in a rather thick layer which forms the outer wall of the oocyst. The basophilic granules, which stain blue with eosin-methylene blue, follow the eosinophilic GENUS Eimeria SCHNEIDER, 1875 123 granules to the periphery of the gamete, where they flatten out and form a thinner layer inside the outer one. A third group of granules arises within the cytoplasm later; they resemble the plastic granules and migrate toward the periphery but do not reach it. Roudabush (1937) be- lieved that they might take part in the formation of the sporocyst walls. Roudabush (1937) did not observe fertilization. He found micro- gametes in the tissues around the macrogametes, but was uncertain whether fertilization instigated oocyst wall formation. Roudabush (1937) calculated that E. nieschulzi could produce 460,800 to 4,819,200 oocysts per oocyst fed. Hall’s (1934) data showed that the total numbers of oocysts actually produced per oocyst fed were 62,000 when | oocyst was fed; 1,455,000 when 6 oocysts were fed; 1,389,000 when 15 oocysts were fed; 1,098,000 when 75 oocysts were fed; 1,029,666 when 150 oocysts were fed; and 144,150 when 2,000 oocysts were fed. According to Hall (1934) and Becker (1934), oocysts continue to be discharged for five to six days after a single inoculation, and for five to eight days after repeated inoculations. Prepatent Period: Seven to eight days. Sporogony: Dieben (1924) described sporogony in the developing oocysts; he studied living, unstained material. After about half an hour the protoplasm of the new oocyst has contracted into a globule, the spo- ront. After a little more than one day, a clear line appears in the sporont; it extends lengthwise through the whole sporont. Dieben (1924) thought this might indicate “chromosomes-conjugation” after fertilization. Very often a little body appears to leave the sporont; Dieben (1924) believed that this might indicate reduction division. The sporont then becomes irregular, and bright granules appear at four corners. The sporont becomes rectangular, growing into the so-called “Buckel” or “stringed” stage. The grooves between the corners deepen so that four daughter- globules are formed, each containing a bright granule. The latter forms the top of a transparent pyramid which appears on each globule. These pyramids diminish a little, forming a “pseudopyramid” stage. ‘They then increase in size, forming the pyramid stage. Four rounded sporoblasts and an oocyst residuum are then formed. The sporoblasts become ovoid and form sporocysts, each containing two sporozoites and a residuum. Landers (1960) attempted to induce excystation in vitro by treating sporulated oocysts with pepsin, trypsin, pancreatin, pancreatic lipase, beef, dog, and rat bile, leuco methylene blue-methylene blue, ethanol- acetaldehyde, 30 pounds air pressure, vacuum (—28 inches of mercury), and fresh blood, but without success. However, the oocysts excysted and produced typical infections following injection directly into the skeletal muscle or peritoneal cavity. 124 THE COCCIDIAN PARASITES OF RODENTS Type Host: Rattus norvegicus (Norway rat). Other Hosts: Rattus rattus (black rat), Rattus hawatiensis (found in the present study). Location: Small intestine, particularly the middle part thereof. Ac- cording to Roudabush (1937), E. nieschulz occurs in the epithelial cells along the base of the villi and in the glands, but usually not as far as the fundi of the glands or in the Paneth cells. The schizonts are usually above the host cell nuclei, while the great majority of the macrogametes are beneath them. (According to Matubayasi, 1938, the first stage of the schizont is beneath the nucleus and pushes it up to the surface as it grows.) Geographic Distribution: Worldwide. Pathogenicity: According to Pérard (1926), E. nieschulzi may produce severe diarrhea or even death in young rats less than six months old. Becker (1934) found that the size of the infecting dose determines the severity of the symptoms. If a rat is fed up to 2,000 oocysts daily for five days, it will become immune without developing clinical evidence of severe disease. However, 30,000 to 100,000 oocysts will produce severe diarrhea on about the seventh day and may cause death. Becker and his associates studied the effects of various constituents of the ration on oocyst production by E. nieschulz (Becker, 1939, 1941, 1942; Becker and Dilworth, 1941; Becker, Manresa, and Smith, 1943; Becker and Smith, 1942). They found, among other things, that thiamin and riboflavin decreased oocyst production, while pantothenic acid increased it and nicotinic acid had no effect. Cross-Transmission Studies: Dieben (1924) transmitted this species from Rattus norvegicus to R. rattus and vice versa, but was unable to transmit it to the house mouse, guinea pig, or domestic rabbit. Pérard (1926) was unable to infect the house mouse or domestic rabbit with £. nieschulzi. Prevalence: Dieben (1924) reported that all the young wild R. norvegicus and R. rattus he examined in the Netherlands were infected. Pérard (1926) noted that this organism caused considerable loss among rats in the Pasteur Institute in Paris. However, Becker (1934) found it in only 1 of 12 wild R. norvegicus near Ames, Iowa, and did not find it in 500 to 600 laboratory rats at Iowa State College. Beltran and Pérez (1950) found it in | per cent of 200 wild R. norvegicus in Mexico City. We found E. nieschulzi in the present study of | out of 9 R. hawaziensis from the vicinity of Honokaa on the island of Hawaii and in 3 out of 11 R. norvegicus from farms in the vicinity of Sullivan, Illinois. Bonfante, Faust, and Giraldo (1961) reported finding FE. nieschulzi in 66 per cent GENUS Eimeria SCHNEIDER, 1875 125 of 71 wild R. norvegicus and 69 per cent of 16 wild R. rattus in Cali, Colombia. Remarks: The confusion regarding the correct specific names for the coccidia of Rattus norvegicus has been discussed under E. mvyairit (p17). Yakimoff (1935) described a coccidium from the Norway rat in Russia under the name EF. halli. Its oocysts were ovoid, subspherical, or spherical, with a smooth, yellowish wall, no micropyle, no oocyst residuum or polar granule, but with a sporocyst residuum. The ovoid forms measured 18-32 x 13-22 u, with a mean of 24.0 17.2 u, the subspherical forms measured 14-27 x 12-24 u, and the spherical forms measured 18-20 u. Yakimoff and Gousseff (1936a) reported this species from a black rat in White Russia. This form does not differ significantly from E. nieschulzi. Roudabush (1937) was dubious about its validity, and we consider it a synonym. Becker and Hall (1933) found that there was no cross-immunity be- tween FE. nieschulzi and E. separata. Rats which had developed immunity to one species as the result of previous infection were not immune to the other. EIMERIA SEPARATA BECKER AND HALL, 1931 (Plate 25, Figs. 188-193; Plate 27, Figs. 215-227; Plate 38, Figs. 326-332) Eimeria separata Becker and Hall, 1931: 131; Becker, Hall, and Hager, 1932: 299-316; Becker, 1934; Yakimoff and Gousseff, 1935: 261-265; Yakimoff and Gousseff, 1936a: 504-508; Roudabush, 1937: 135-163; Matubayasi, 1938: 144— 163; Beltran and Pérez, 1950: 71-78; RySavy, 1954: 131-174; Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts predominantly ellipsoidal, although subspherical and ovoid forms may also occur. Oocyst wall smooth, colorless to pale yellowish, composed of a single layer about 0.6 u thick. The oocyst wall was illustrated by Becker, Hall, and Hager (1932) as apparently com- posed of two layers, but this was probably due to the manner of drawing; they did not mention the number of layers in their description, and we found only a single layer in the forms from R. norvegicus which we studied. Oocysts 13-19 x 11-17 u, with a mean of 16.1 x 13.8 u and a mean length-width ratio of 1.16, according to Becker, Hall, and Hager (1932). Ninety oocysts from three R. norvegicus from Hlinois measured by us in the present study were 10-16 x 10-14 u, with a mean of 13.411.6 u; their length-width ratios ranged from 1.0-1.4, with a mean of 1.15. Micropyle absent. One to three oocyst polar granules present. Oocyst residuum absent. Sporocysts ellipsoidal, with a tiny Stieda body. Twelve sporocysts from R. norvegicus from Illinois meas- 126 THE COCCIDIAN PARASITES OF RODENTS ured by us in the present study were 8-105-6 u, with a mean of 9.0x 5.1 u; their length-width ratios ranged from 1.6—-1.9, with a mean of 1.76. Sporocyst wall very thin. Sporozoites elongate, often with a clear globule at one end, lying lengthwise, head to tail, in the sporocysts. Sporocyst residuum small and compact, sometimes not discernible. Sporulation Time: Less than 36 hours (three days at room temperature in Liberia for the strain from R. defua). The following description was given by Levine, Bray, Ivens, and Gunders (1959) of the form they found in Rattus (Dephomys) defua in Liberia: Oocysts subspherical, ellipsoidal, or slightly ovoid. Oocyst wall smooth, colorless to pale yellowish or greenish, composed of a single layer about | uw thick. Micropyle absent. “Twenty sporulated oocysts measured 16-21 x 15-17 u, with a mean of 18.4 15.8 u; their length- width ratios ranged from 1.1—1.3, with a mean of 1.16. Oocyst polar granule present. Oocyst residuum absent. Oocyst wall collapses rather quickly in Sheather’s sugar solution. Sporocysts 11 x 7 u, almost ellipsoidal, with a small Stieda body. Sporocyst wall thin. Sporocyst residuum present, small. Some sporozoites appear to have a clear globule at one end. Sporozoites lie lengthwise in sporocysts but are curled at the ends. The form found in Rattus hawaiiensis in the present study had the following characteristics: Oocysts ellipsoidal to subspherical. Oocyst wall smooth, pale yellowish, composed of a single layer 0.6-0.8 u thick. Micropyle absent. Thirty sporulated oocysts measured 12-16 x 10-13 u, with a mean of 13.5x11.6 u; their length-width ratios ranged from ].1-1.3, with a mean of 1.16. One to two oocyst polar granules present. Oocyst residuum absent. Sporocysts almost ellipsoidal, with a small, some- what flat Stieda body. Four sporocysts measured 8-9 x 5 u, with a mean of 8.8x5.1 u; their length-width ratios ranged from 1.6-1.7, with a mean of 1.68. Sporocyst residuum small to very small or absent. Sporo- zoites oriented longitudinally, head to tail, in sporocysts, with a pale yellowish globule at the large end. Schizogony: Roudabush (1937) described the endogenous stages of F. separata in detail. Sporozoites are found in the lumen of the cecum as early as six hours after infection, but some may also be found as long as three days after infection. They measure 8-10 x 2-3 u, with a mean of 9.5 x 2.3 u. The nucleus is central and has a central karyosome; there is a siderophilic, refractile globule at either end of the sporozoite. The sporozoite enters an intestinal cell and rounds up to form a first-generation schizont. This divides by multiple fission to form 6 to 12 (mean, 8) first-generation merozoites; there is no residuum. These merozoites measure 11—13 x 2—3 u, with a mean of 11.9x 2.2 u, and have a central nucleus with a central karyosome and granular cytoplasm with- GENUS Eimeria SCHNEIDER, 1875 127 out refractile globules. They break out of the host cell in one day, enter new host cells, round up and form second-generation schizonts. Each of these produces 4 to 6 (mean, 5.5) second-generation short, broad merozoites measuring 6-9 x 2-3 u, with a mean of 7.7 x 2.3 u. These break out of the host cell in two days after infection and enter new host cells, where they round up to form third-generation schizonts. Each third-generation schizont forms 2 to 6 (mean, 4) third-generation mero- zoites by the third day after infection. They measure 13-15x2-3 u, with a mean of 13.6 2.6 u. No residuum is left. The third-generation merozoites differ from the previous two not only in size but also in the fact that their anterior end stains an intense red with eosin-methylene blue or brownish yellow with iron hematoxylin. Gametogony: The third-generation merozoites enter new host cells. Roudabush (1937) considered that they formed microgametocytes and macrogametocytes, but the cells he considered macrogametocytes are actually macrogametes. The young “gametocytes” are indistinguishable and have an oval vacuolated area which contains an eccentric chromatin mass. The microgametocyte nucleus divides repeatedly, and the nuclei thereby produced migrate to the periphery of the cell. Here the micro- gametes form; they are 2—3 u long and less than 0.5 uw wide at the anterior end, and taper to a point posteriorly; they have two flagella. The macrogametes contain both eosinophilic plastic granules and hematoxylinophilic granules. These pass to the periphery of the cell and flatten out, the eosinophilic granules forming the outer oocyst wall and the hematoxylinophilic granules the thin inner membrane of the oocyst. Meanwhile, a second set of plastic granules forms. ‘They migrate to just within the inner wall, where they still remain when the oocyst leaves the host cell; their further development has not been followed. Roudabush (1937) believed that fertilization takes place in the tissues during the early oocyst wall formation. Mature oocysts appear five to six days after infection. Roudabush (1937) calculated that E. separata could produce 384 to to 3,456 (mean, 1,536) gametocytes per oocyst fed. The numbers of oocysts produced would be these figures minus the numbers of microgametocytes. According to Becker, Hall, and Hager (1932), oocysts continue to be discharged for three to four days after a single inoculation, and from four to six days after repeated inoculations. The average number of oocysts discharged during the patent period following repeated inocula- tions was 2.46 x 10°. Prepatent Period: Five to six days. Type Host: Rattus norvegicus (Norway rat). Other Hosts: Rattus rattus (?). Matubayasi (1938) found this species 128 THE COCCIDIAN PARASITES OF RODENTS “often” in wild rats (R. norvegicus and R. rattus) which he examined in Japan. However, he did not specify whether it occurred in one or both host species. Rattus (Dephomys) defua. Levine, Bray, Ivens, and Gunders (1959) reported E. separata from this species in Harbel, Liberia. Rattus hawatriensis. Found in the present study. Location: Cecum and colon. According to Roudabush (1937), infec- tion is heavier in the cecum. E. separata occurs exclusively in the cells of the surface epithelium, and not in the epithelium of the glands. The schizonts, gametocytes, and gametes are found below the host cell nuclei. Roudabush found some stages, especially the macrogametes, below the epithelial layer, but thought this may have been due to the plane in which the sections were cut. A similar appearance was noted by Matu- bayasi (1938). Geographic Distribution: North America (Iowa, Illinois, Mexico), Hawaii, Europe (Russia, Czechoslovakia), Japan, Africa (Liberia). Pathogenicity: According to Becker (1934), E. separata is not patho- genic enough to cause clinical symptoms even when heavy infective doses are administered. However, some hyperemia of the cecal and colon walls was noted when the rats were killed on about the sixth day of infection. Cross-Transmission Studies: None. Prevalence: Beltran and Pérez (1950) found this species in 11 per cent of 200 wild R. norvegicus in Mexico City. In the present study, we found it in 10 of 11 wild R. norvegicus from farms in the vicinity of Sullivan, Illinois. Levine, Bray, Ivens, and Gunders (1959) found it in the only Rattus defua which they examined from Harbel, Liberia. It was found in the present study in 2 out of 9 R. hawariensis from the vicinity of Honokaa on the island of Hawaii. Remarks: Becker and Hall (1933) found that there was no cross-im- munity between EL. separata and E. nieschulzi. Rats which had developed immunity to one species as the result of previous infection were not immune to the other. EIMERIA HASEI YAKIMOFF AND GOUSSEFF, 1936 (Plate 38, Figs. 333 and 334) Eimeria hasei Yakimoft and Gousseff, 1936a: 504-508. Description: Oocysts ovoid, ellipsoidal, or spherical. Oocyst wall smooth, illustrated as composed of a single layer. Spherical oocysts 12-24 u in diameter, with a mean of 16.1 u. Other oocysts 16-20 12-17 u, with a length-width ratio of 1.1-1.4. Micropyle absent. Oocyst polar granule present. Oocyst and sporocyst residua absent. Sporocysts 8.55 u. GENUS Eimeria SCHNEIDER, 1875 129 Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Rattus rattus (black rat). Location: Intestine. Geographic Distribution: USSR (White Russia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoft and Gousseff (1936a) found this species in 2 per cent of 47 black rats in White Russia. EIMERIA NOCHTI YAKIMOFF AND GOUSSEFF, 1936 (Plate 39, Fig. 335) Eimeria nochti Yakimoft and Gousseff, 1936a: 504-508; RySavy, 1954: 131- Nia (=). Description: Oocysts ovoid. Oocyst wall smooth, described by Yakimoff and Gousseff (1936) as “doppelkonturige,” but illustrated as composed of a single layer. Oocysts 15-24 x 12-22 u, with a mean of 17.2x 14.2 u. Oocyst length-width ratio 1.1-1.4, with a mean of 1.22. Micropyle absent. Oocyst polar granule absent. Oocyst and sporocyst residua absent. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Rattus rattus (black rat). Other Host: Rattus norvegicus (Norway rat) (?). Location: Intestine. Geographic Distribution: USSR (White Russia), Europe (Czechoslo- vakia?). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseff (1936a) found this species in 2 per cent of 47 black rats in White Russia. Remarks: RySavy (1954) reported E. nochti from Rattus norvegicus in Czechoslovakia. The oocysts of his form measured 14—25 x 13-20 u, with a mean of 19.4% 13.8 uw. However, they differed from both E. nochti and the other coccidia reported from Rattus in shape, being pointed at both ends. EIMERIA RATTI YAKIMOFF AND GOUSSEFF, 1936 (Plate 39, Fig. 336) Eimeria ratti Yakimoff and Gousseff, 1936a: 504—508. Description: Oocysts cylindrical to ovoid. Oocyst wall smooth, de- scribed by Yakimoff and Gousseff (1936a) as “doppelkonturige,” but illustrated as composed of a single layer. Oocysts 16-28 x 15-16 u, with 130 THE COCCIDIAN PARASITES OF RODENTS a mean of 2315 u. Oocyst length-width ratio 1.1-1.9, with a mean of 1.61. Micropyle absent. Oocyst residuum absent. Oocyst polar granule present. Sporocyst residuum present. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Rattus rattus (black rat). Location: Intestine. Geographic Distribution: USSR (White Russia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseff (1936a) found this species in 4 per cent of 47 black rats in White Russia. Remarks: This species appears to differ from E. separata Becker and Hall, 1931 only in size and in the fact that some of its oocysts are cylindrical. However, only ten oocysts were measured. Future work may indicate that E. ratti is a synonym of EF. separata. EIMERIA PRAOMYSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 * (Plate 28, Fig. 228) Eimeria praomysis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts subspherical to ellipsoidal. Oocyst wall pale yellowish to brownish, smooth to somewhat rough, composed of a single layer (confirmed by breaking oocyst) approximately | u thick. Micropyle absent. Thirty-one oocysts measured 17-24 x 16-23 u, with a mean of 20.6 x 19.1 uw; their length-width ratios ranged from 1.0-1.3, with a mean of 1.08. One or two oocyst polar granules present. Oocyst residuum absent. Sporocysts ellipsoidal to slightly ovoid, with Stieda body. Six sporocysts measured 10-12 6-7 u, with a mean of 11.2x6.5 u; their length-width ratios ranged from 1.6-1.8, with a mean of 1.72. Sporocyst residual granules present, rather coarse. Sporozoites lie more or less lengthwise in sporocysts. Sporulation Time: Six days at room temperature in Liberia. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Rattus (Praomys) tullbergi rostratus. Location: Intestinal contents. Geographic Distribution: Liberia. Pathogenicity: Unknown. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. GENUS Eimeria SCHNEIDER, 1875 131 Cross-Transmission Studies: None. Prevalence: This species was found in one of three rats examined from Harbel, Liberia. EIMERIA SP. FANTHAM, 1926 Eimeria sp. Fantham, 1926: 560-570. Description: Oocysts oval, 16-21 x 15-16 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Rattus (Mastomys) coucha (syn., Mus coucha) (multimam- mate mouse). Location: Chiefly jejunum and ileum. Geographic Distribution: South Africa. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. Remarks: Fantham (1926) stated that this species might be a variety of Eimeria falciformis, but this is unlikely. EIMERIA FALCIFORMIS (EIMER, 1870) SCHNEIDER, 1875 (Plate 28, Figs. 229-240; Plate 39, Figs. 337-339) Gregarina falciformis Eimer, 1870. Eimeria falciformis (Eimer, 1870) Schneider, 1875: xl-xlv; Wenyon, 1926; Yakimoff, Gousseff, and Suz’ko, 1945: 172-183: Koffman, 1946: 424-442; Cordero del Campillo, 1959: 351-368; Cerna, 1962: 1-13 (?). Coccidium falciforme (Eimer, 1870) Schuberg, 1892. Gregarina muris Rivolta, 1878: 220-235. Pfeifferia schubergi (Labbé, 1896) Labbé, 1899. Eimeria schubergi (Labbé, 1896) Doflein, 1916. Eimeria-like species. Clarke, 1895: 277-283. [non] Eimeria schubergi (Schaudinn, 1900) [non] Eimeria falciformis: Elton et al., 1931: 657-721 [see Eimeria sp. (RySavy, 1954)—p. 113]; RySavy, 1954: 131-174 (pro parte) [see Eimeria sp. (RySavy, 1954) —pp. 81 and 113]; Cerna and Daniel, 1956: 19-23 [see Eimeria sp. (RySavy, 1954)—p. 113]; Ring, 1959: 381-401. Description: Oocysts broadly ovoid to subspherical or spherical, 16-21 x 11-17 u. Oocyst wall smooth, apparently colorless. Muicropyle absent. Oocyst residuum either absent or present. Sporocyst Stieda body present, although not always illustrated. Sporocyst residuum present. Sporozoites lie longitudinally in sporocysts. The above description is based on the earlier literature. A more com- plete description, based on a laboratory strain obtained from Weybridge, England, was given by Cordero del Campillo (1959). Unfortunately, he gave no drawing but only a rather blurred photomicrograph. According 132 THE COCCIDIAN PARASITES OF RODENTS to his description, the oocysts are ellipsoidal, with a smooth, colorless wall without a micropyle. He made a biometric study of the dimensions of 1,527 oocysts during a 20-day period. ‘They ranged from 15-26 x 13-24 u; their means varied on different days from 19-22 x 16-19 u, and the mean of an “ideal” oocyst was considered to be 21x18 u. The oocysts tended to increase in size from the first to the fifth or sixth days of the patent period and then decreased in size. The oocyst length-width ratio ranged from 1.1—1.2, with a mean of 1.17. There were one or two cylindrical oocyst polar granules measuring 31.5 uy, but no oocyst residuum. The sporocysts measured 10—12 x 6-8 u and had a conspicuous Stieda body (which he called a micropyle). The sporocyst residuum was subspherical and measured 4—6 x 3-5 u. The sporozoites had a large, refractile globule 3 uw in diameter. The following description is of oocysts found in the present study in a Mus musculus from Hlinois and assigned to this species: Oocysts sub- spherical to spherical. Oocyst wall smooth, almost colorless to pale yellowish, composed of a single layer about | u thick. Eight sporulated oocysts measured 15-18 x 13-16 u, with a mean of 16.7 14.6 u; their length-width ratios ranged from 1.1—1.2, with a mean of 1.14. Micropyle absent. Oocyst polar granule absent. Oocyst residuum absent. Sporocysts elongate ovoid, about 117 u, thin-walled, with a tiny Stieda body. Sporocyst residuum present, usually compact, finely granular. Sporozoites with a large, homogeneous yellowish globule at the large end. Sporozoites lie lengthwise in sporocysts. In addition to the above, found in the same mouse was about an equal number of oocysts which differed from the above description only in having an oocyst polar granule. ‘Ten of these oocysts measured 14-17 x 12-16 u, with a mean of 16.1 x 14.1 u; their length-width ratios ranged from 1.0-1.2, with a mean of 1.15. Although the older descriptions of E. falciformis omit mention of an oocyst polar granule, Cordero del Campillo (1959) found one. Some earlier authors mentioned an oocyst residuum and it is possible that they used this term instead of polar granule. That a polar granule may be formed and then disappear later on in the course of sporogony may account for the difference we ob- served. All our material was used up in making these observations. Clarification of this point must await further studies, but in the mean- time it is considered best not to assign our two forms to different species. Sporulation Time: Five to six days according to Wenyon (1926), three days according to Nieschulz and Bos (1931), Koffman (1946), and Cordero del Campillo (1959). Schizogony: Although schizogony has not been studied in detail, some stages have been described by Eimer (1870), Schneider (1875), Schuberg GENUS Eimeria SCHNEIDER, 1875 133 (1892), (1895), Clarke (1895), Reich (1913), Reichenow (1921), Reimer (1923), Wenyon (1926), Kiedrowski (1925), and Nieschulz and Bos (1931). The protozoa are found in the epithelial cells of the small intestine, but they may also occur in the stomach and large intestine. The generations of merozoites have not been differentiated. The mature schizonts may produce from 8 or possibly fewer to 30 or more elongate, falciform merozoites which often lie around a central residual mass of cytoplasm like the ribs of a barrel; they may also spiral somewhat. Merozoites appear in the feces three days after experimental infection and persist until the twelfth day, being most numerous on the fourth day after infection. Gametogony: Gametogony has been described particularly by Reich (1913) and also by Wenyon (1926). The microgametocytes are large, irregularly shaped cells on the surface of which numerous nuclei develop. Each nucleus gives rise to a comma-shaped biflagellate microgamete. The macrogametes are ovoid to subspherical and contain a central nucleus and numerous plastic granules. These coalesce to form the oocyst wall, but leave a micropyle at one end. Fertilization then takes place according to Reich (1913), after which the micropyle disappears. The oocysts then pass out of the body. Prepatent Period: Oocysts first appear in the feces on the fourth day after infection according to Reimer (1923) or on the fifth day according to Nieschulz and Bos (1931) and Cordero del Campillo (1959). According to Nieschulz and Bos, they reach their maximum between the sixth and twelfth days, and by the fifteenth day half the experimentally in- fected mice are no longer shedding oocysts. Type Host: Mus musculus (house mouse). Location: Epithelial cells of the small and large intestines. Clarke (1895) and Reich (1913) reported it also from the stomach, but Becker (1934) considered that it remained to be established whether the stomach forms were Eimeria rather than Cryptosporidium muris. According to Reimer (1923), the parasites are found in the crypts but not in the tips of the villi of the small intestine and in the mucosa of the large intestine. Geographic Distribution: Worldwide. Pathogenicity: Light infections have little effect, but heavy ones may cause diarrhea and even death. A catarrhal enteritis may occur, together with desquamation of the intestinal epithelium and hemorrhage. The feces may contain blood. According to Nieschulz and Bos (1931), mor- tality is highest from the fourth to the eighth day after infection. Cordero del Campillo (1959) saw no clinical signs in mice infected with 55,000 oocysts; mice infected with 100,000 to 500,000 oocysts lost their appetites; mice infected with 2 million oocysts had dysentery and hemorrhages in the small intestine. 134 THE COCCIDIAN PARASITES OF RODENTS Cross-Transmission Studies: Noller (1920) was unable to infect the rat or dog with E. falciformis, and Pérard (1926) was unable to infect the rat with it. Prevalence: E. falciformis is apparently much commoner in Europe than in the United States. Wenyon (1926) stated that it was common in England, and Yakimoff, Gousseff, and Suz’ko (1945) found it in 3 out of 18 wild M. musculus severtzovi in Vadzhikistan. On the other hand, Becker (1934) saw it only once in a laboratory mouse, and examined nearly 50 wild house mice in Jowa without finding any coccidia. We found it in 1 out of 42 wild M. musculus captured on farms in the vicinity of Sullivan, Illinois, but we failed to find it in several laboratory mouse colonies and in half a dozen wild house mice captured in Urbana, Illinois. Bonfante, Faust, and Giraldo (1961) reported finding E£. falci- formis in all of the 5 wild Mus musculus they examined in Cali, Colombia. Remarks: RySavy (1954) reported E. falciformis from Mus musculus, Apodemus flavicollis, A. sylvaticus, Microtus arvalis, and Clethrionomys glareolus in Czechoslovakia. The description he gave was similar to that of E. falciformis, but he did not describe the forms from each host separately. For the reasons given in the discussion of these forms in Microtus arvalis (p. 81), it is very doubtful that the forms RySavy saw in the hosts other than M. musculus were E. falciformis. However, since he did not describe them separately from each host, it is useless to attempt to assign names to them. Cerna and Daniel (1956) reported E. falciformis from Apodemus flavicollis and Clethrionomys glareolus in Czechoslovakia. They gave the same dimensions as RySavy, but mentioned that the oocyst wall was more than | u thick. For the reasons given above, these forms were undoubtedly not E. falciformis either, but cannot be assigned separate names. Cerna (1962) reported finding E. falciformis not only in Mus musculus but also in Apodemus flavicollis and A. sylvaticus. She studied the coccidia in experimentally infected M. musculus, but her description differed in some ways from the descriptions of all other coccidia from this host, and it is uncertain whether she was dealing with a pure infec- tion. Hence, the information in her paper is summarized below, but without attempting to establish the species involved. Most of the oocysts were broadly “oval,” 14-23 x 14-20 u; often slightly narrowed toward one or both poles. ‘They sporulated in two to three days at room tem- perature in 1.5%, potassium dichromate solution. ‘The sporocysts meas- ured 4—5 x 3-4 uw (according to Cerna’s text) or 7-8 x 4.5 uw (according to her table). A small polar body was occasionally (according to Cerna’s text) or usually (according to her table) present. She found endogenous GENUS Eimeria SCHNEIDER, 1875 135 stages only in the mucosa and submucosa of the colon and cecum, and never in the small intestine. She described two schizogonic stages. The first schizonts appeared as early as the second day after infection, measured 4—7 x 4-6 u, and gave rise to only 4 merozoites. The second- generation schizonts measured 19-25 x 13-18 u and produced 12 to 16 merozoites. The merozoites were slender, 9-10 x 2-3 u, slightly narrowed at both ends, with an excentric nucleus. The mature microgametocytes measured 11-16 6-11 u, and the mature macrogametes measured 12- 18x 12-15 u. Elton et al. (1931) and Ring (1959) reported finding Eimeria falciformis in Apodemus sylvaticus, Clethrionymus glareolus, and Microtus agrestis hirtus in England. They did not describe the coccidia, which were un- doubtedly not EF. falciformis but species characteristic of their several hosts. EIMERIA HINDLEI YAKIMOFF AND GOUSSEFF, 1938 (Plate 37, Fig. 298) Eimeria hindlei Yakimoft and Gousseff, 1938: 1-3; Svanbaev, 1956: 180-191 (from Mus musculus). [non] Eimeria hindlei: Svanbaev, 1956: 180-191 (from Apodemus sylvaticus— see E. prasadi); RySavy, 1954: 131-174 (see E. rysavyi); Cerna, 1962: 1-13 (see E. rysavyt). Description: Oocysts ovoid, smooth, 22-27 x 18-21 u (28.4 x 22.0 u and greenish in color according to Svanbaev, 1956). Oocyst length-width ratio 1.2—-1.4. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid; according to Svanbaev (1956) they measure 8.6 x 6.4 u. Sporozoites illustrated with a clear globule at one end. The situation regarding the sporocyst residuum requires discussion. In the paper in which they described E. hindlei, Yakimoff and Gousseft (1938) also described four other species of Eimeria from Mus musculus: E. keilint, E. krijgsmanni, E. musculi, and E. schueffnert. Their state- ments regarding the sporocyst residua of these five species were respec- tively: “We could not find residual bodies in the sporocysts,” “We could not distinguish any residual bodies in the sporocysts,” ““We were unable to discover whether the sporocysts had residual bodies,” and (for the last two) “The presence of a residual body in the sporocysts was uncer- tain.” They showed no residual bodies in the rough sketches which illustrated their paper. In his description of FE. hindlei from Mus musculus, Svanbaev (1956) stated that a sporocyst residual body was present. The question arises whether the E. hindlei seen and named by Yakimoff and Gousseff (1938) actually lacked a sporocyst residuum or whether they meant by their statements that they were unable to deter- mine whether one was present or not. If the former is the case, Svanbaev’s 136 THE COCCIDIAN PARASITES OF RODENTS form should not be assigned to this species. If the latter is the case, it could be. Until this point has been investigated further, it is considered better to assign them to the same species. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Feces. Geographic Distribution: USSR (White Russia, western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimofft and Gousseft (1938) found E. hindlei in 1 of 20 mice examined in White Russia. Remarks: Svanbaev (1956) described an Eimeria under the name E. hindlei from the field mouse, Apodemus sylvaticus, in western Kazakh- stan. It did not belong to this species, and we have named it Eimeria prasadi (see p. 110). Rygavy (1954) described an Eimeria under the name E. hindlei from Apodemus sylvaticus and Clethrionomys glareolus in Czechoslovakia. He did not differentiate between the forms from the two hosts, the oocysts failed to sporulate, and RySavy made no mention of Galli-Valerio’s or Pellérdy’s work cited on page 110. On the basis of the meager mor- phological information supplied, this form(s?) does not disagree with the description of E. hindlei, but in view of the failure of Pellérdy’s cross- transmission attempts, it is unlikely that it is this species; as a matter of fact, Ry’avy was probably dealing with a different species in each host. The form from Apodemus sylvaticus agrees most closely with E. prasadi from the same host, but under the circumstances it is considered best not to assign it to any species. RySavy (1957) gave further information regarding the form from C. glareolus, and Cerna (1962) described its oocysts and sexual stages. We have named this form Eimeria rysavyi (see page 82). EIMERIA KEILINI YAKIMOFF AND GOUSSEFF, 1938 (Plate 40, Fig. 347) Eimeria keilini Yakimoft and Gousseff, 1938: 1-3. [non] Eimeria keilini: RySavy, 1954: 131-174 [see Eimeria sp. (RySavy, 1954)— p- 113]; Cerna and Daniel, 1956: 19-23 [see Eimeria sp. (RySavy, 1954)—p. 113); Cerna, 1962: 1-13 [see Eimeria sp. (RySavy, 1954)—p. 113]. Description: Oocysts smooth, yellowish, pointed at both ends. Micro- pyle absent. Oocysts 24-32 x 18-21 u, with a mean of 28.8 x 19.4 u. Oocyst length-width ratio 1.2-1.7, with a mean of 1.54. Oocyst polar granule and oocyst residuum absent. Sporocysts 12.2 6.1 u. The authors stated, “We GENUS Eimeria SCHNEIDER, 1875 137 could not distinguish any residual bodies in the sporocysts.” See the description of E. hindlei (p. 135), for a discussion of the meaning of this statement. Stieda body not mentioned and not shown in the rather sketchy illustration. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Feces. Geographic Distribution: USSR (White Russia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseft (1938) found E. keilint in 1 of 20 mice examined in White Russia. Remarks: Rysavy (1954) described a coccidium from Apodemus sylvati- cus in Czechoslovakia under the name Eimeria keilini. While the shape of its oocysts as shown in his illustration resembled that of E. kezlini, the oocysts did not sporulate. Cerna and Daniel (1956) and Gerna (1962) found what was presumably the same form in A. flavicollis in Czechoslo- vakia. It sporulated, and its sporocysts were smaller than those of FE. keilini, measuring 8-9 x 6-7 u. In addition, its wall was colorless. In view of these differences, of Pellérdy’s (1954) failure to transmit four species of Eimeria from Apodemus flavicollis to Mus musculus, and of the failure of various attempts to transmit Eimeria between Mus and Rattus, it is doubtful whether RySavy’s form is E. keilini. Its shape is unlike that of any of the six species of Eimeria so far described from Apodemus. In the absence of more information, it is discussed on page 113 as Eimeria sp. (RySavy, 1954). EIMERIA KRIJGSMANNI YAKIMOFF AND GOUSSEFF, 1938 (Plate 42, Fig. 376) Eimeria krijgsmanni Yakimoft and Gousseff, 1938: 1-3. Eimeria kriygsmanni [sic]: Svanbaev, 1956: 180-191 (from Mus musculus). [non] Eimeria krijgsmani [sic]: RySavy, 1954: 131-174 (see E. svanbaevi). [non] Eimeria kriygsmanni [sic]: Svanbaev, 1956: 180-191 (from Apodemus sylvaticus—see E. suanbaevi); Svanbaev, 1958: 183-186 (from Ochotona pallas?); Svanbaev, 1962: 23-39 (see E. peschankae). Description: Oocysts described as “‘oval,’”’ but illustrated as ellipsoidal, smooth, colorless or yellowish. Micropyle absent. Oocysts 18-23 x 13-16 u, with a mean of 21.9 14.8 u (25.4 18.8 u, according to Svanbaev, 1956). Oocyst length-width ratio 1.2-1.6, with a mean of 1.37 (1.35, according to Svanbaev, 1956). Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, 12.3x8.7 u according to Svanbaev (1956). Sporocyst residuum absent according to Svanbaev (1956). (Yakimoff and 138 THE COCCIDIAN PARASITES OF RODENTS Gousseff (1938) stated, “We were unable to discover whether the sporocysts had residual bodies.”) Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Feces. Geographic Distribution: USSR (White Russia, western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseff (1938) found this species in 7 of 20 mice in White Russia. Remarks: Svanbaev (1956) described an Eimeria under the name “FE. kriygsmanni” (lapsus for E. krijgsmanni) from Apodemus sylvaticus in western Kazakhstan. It did not belong to this species, and we have named it Eimeria svanbaevi (see p. 111). Svanbaev (1958) described an Eimeria under the name “E. kriygs- manni” (lapsus for E. krijgsmanni) from the Mongolian pika, Ochotona pallasi, in Kazakhstan. However, since Ochotona is a lagomorph rather than a rodent, it is very doubtful that the pika form belongs to this species. Svanbaev (1962) described an Ez:meria under the name “E. kriygsmanni” from the tamarisk gerbil, Meriones tamariscinus, in Kazakhstan. How- ever, it did not belong to this species, and we have named it Eimeria peschankae (see p. 89). RySavy (1954) described an Eimeria under the name “E. krijgsmani” (lapsus for E. krijgsmanni) from Apodemus sylvaticus in Czechoslovakia. He, too, was unaware of Galli-Valerio’s (1932, 1940) and Pellérdy’s (1954) work. His description was so incomplete that it is not certain what species he was dealing with. We are tentatively assigning it to Eimeria svanbaevi (Os JANN). Gerna (1962) considered E. krijgsmanni to be a synonym of E. schueff- neri; however, her reasons were inadequate and based only on disagree- ment with Yakimoff and Gousseff’s interpretation. EIMERIA MUSCULI YAKIMOFF AND GOUSSEFF, 1938 (Plate 37, Fig. 299) Eimeria musculi Yakimoff and Gousseff, 1938: 1-3; Svanbaev, 1956: 180-191 (from Mus musculus). [non] Eimeria musculi: Svanbaev, 1956: Ibid. (from Apodemus sylvaticus—see E. russiensis) (from Microtus arvalis—see E. arvicolae) (from Mertones tamari- scinus—see E. tamariscini); Svanbaev, 1958: 183-186 (from Ochotona pallasi?). Description: Oocysts spherical, smooth, greenish, 21-26 u in diameter. Micropyle absent. Polar granule absent. Oocyst residuum absent. Sporo- GENUS Eimeria SCHNEIDER, 1875 139 cysts broadly ovoid (109 u according to Svanbaev, 1956). Sporocyst re- siduum absent according to Svanbaev (1956). (Yakimoff and Gousseff, 1938 stated, ‘““The presence of a residual body in the sporocysts was un- certain.’’) Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Feces. Geographic Distribution: USSR (White Russia, western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseff (1938) found this species in 1 of 20 mice in White Russia. Remarks: Svanbaev (1956) described coccidia from western Kazakhstan under the name Eimeria musculi not only from Mus musculus but also from Apodemus sylvaticus, Microtus arvalis, and Meriones tamariscinus. (See Remarks under E. prasadi, p. 110, for a discussion of his paper). The form which he described from 4. sylvaticus is E. russiensis (p. 112), that from M. arvalis is E. arvicolae (p. 75), and that from M. tamariscinus is E. tamariscini (p. 88). Svanbaev (1958) described an Eimeria under the name E. musculi from the Mongolian pika, Ochotona pallasi, in Kazakhstan. However, since Ochotona is a lagomorph rather than a rodent, it is very doubtful that the pika form belongs to this species. EIMERIA SCHUEFFNERI YAKIMOFF AND GOUSSEFF, 1938 (Plate 40, Fig. 346) Eimeria schueffneri Yakimoft and Gousseff, 1938: 1-3. [non] Eimeria schueffneri: Cerna, 1962: 1-13 (see E. cernae). Description: Oocysts cylindrical (illustrated with rounded ends), smooth, colorless. Micropyle absent. Oocysts 18-26 x 15-16 u. Oocyst length-width ratio 1.5-1.9. Oocyst polar granule and oocyst residuum absent. Sporocysts ovoid. The authors stated, “The presence of a residual body in the sporocysts was uncertain.” Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Feces. Geographic Distribution: USSR (White Russia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff and Gousseff (1938) found E. schueffneri in 1 out of 20 mice examined in White Russia. 140 THE COCCIDIAN PARASITES OF RODENTS Remarks: Cerna (1962) described a coccidium from Clethrionomys glareolus under the name E. schueffneri. It does not belong to this species, and we are assigning it the name E. cernae (p. 84). EIMERIA HANSONORUM N. SP. (Plate 28, Fig. 241) 2 Description: Oocysts subspherical. Oocyst wall smooth, pale yellowish, composed of a single layer 0.8 uw thick. Fifteen sporulated oocysts measured 15-22 13-19 u, with a mean of 17.9 15.8 u; their length- width ratios ranged from |.1—1.2, with a mean of 1.13. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, thick-walled, with a broad, thick Stieda body. Sporocysts about 9X7 u, with a length-width ratio of about 1.4. Sporocyst residuum composed of loose, coarse granules. Sporozoites pale, lying longitudinally in sporo- cysts. Schizogony and Gametogony: Unknown. No intestinal sections were examined from the host animal. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Intestinal contents. Geographic Distribution: United States (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in | out of 42 M. musculus cap- tured on farms in the vicinity of Sullivan, [linois. Remarks: Seven species of Eimeria have been named from rodents of the genus Mus; six of these have been named from M. musculus. Despite the fact that the descriptions of most of these are incomplete, E. hansonorum appears to differ from all of them. It differs from E. falciformis in the shape of its sporocysts, in the size and shape of its Stieda body, and in lacking clear globules in the sporozoites. It also differs from the classical E. falciformis in having an oocyst polar granule. It differs from E. hindlei in oocyst size and shape. It differs from E. krijgsmanni in oocyst shape. It differs from E. kezlini in oocyst size and shape, in having a polar granule, and in sporocyst size. It differs from E. musculi and E. schueffneri in oocyst shape and in having an oocyst polar granule. In addition, it may differ in sporocyst characters from the last five species, but they were so poorly described and illustrated that it is impossible to determine this. It differs from E. musculoide: in sporocyst shape and in having a prominent Stieda body. This species is named in honor of Dr. Harold C. Hanson, Illinois State Natural History Survey, Urbana, and Dr. Lyle E. Hanson, College of Veterinary Medicine, University of Illinois, Urbana. GENUS Eimeria SCHNEIDER, 1875 141 EIMERIA FERRISI N. SP. (Plate 29, Fig. 242) Description: Oocysts ellipsoidal to subspherical. Oocyst wall smooth, colorless to very pale yellowish, composed of a single layer about 0.9 u thick. Ten sporulated oocysts from a single host animal measured 17-— 20x 14-16 u, with a mean of 17.8 x 15.4 u; their length-width ratios ranged from 1.1-1.2, with a mean of 1.15. Micropyle absent. One to three oocyst polar granules present. Oocyst residuum absent. Sporocysts elongate ovoid, thin-walled, with a small Stieda body. Sporocysts 10-11 x 5-6 u, with a length-width ratio of 1.8-1.9. Sporozoites elongate, lying length- wise head to tail in sporocysts, but bent over at the smaller end. Sporo- zoites with a colorless globule at the large end. Sporocyst residuum absent, or in a few cases represented by a few tiny granules. Schizogony and Gametogony: Macrogametes, microgametocytes, young oocysts and a few schizonts were found in the epithelial cells of the tips of the villi of the cecum. None were in the crypts. All stages were present both above and below the host cell nuclei. The zygotes and young oocysts tended to be more often below than above the host cell nuclei. In some cases the host cells appeared to be displaced somewhat toward the interior of the villus. Macrogametes ranged in size up to 16x 12 u, with a single row of hematoxylin-staining plastic granules. Microgametocytes ranged in size up to 12x 10 u. Oocysts ranged in size up to 17x 13 u. The few schizonts seen measured up to about 129 u and contained about 10 to 16 elongate, curved, sausage-shaped merozoites measuring about 132 u. There was no schizont residuum. Prepatent Period: Unknown. Type Host: Mus musculus (house mouse). Location: Cecum. No endogenous stages were found in the duodenum, jejunum, anterior ileum, or posterior ileum. Geographic Distribution: United States (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in 2 out of 42 M. musculus captured on farms in the vicinity of Sullivan, Illinois. Remarks: Eight species of Eimeria have been named from rodents of the genus Mus; seven of these have been named from M. musculus. Despite the fact that the descriptions of most of these are incomplete, E. ferrisi appears to differ from all of them. It differs from E. falciformis in lacking a sporocyst residuum and from the classical E. falciformis in having an oocyst polar granule. It differs from FE. hindlei in oocyst size and shape. It differs from EF. keilini, E. kriyjgsmanni, E. musculi, E. schueffneri, E. hansonorum, and E. musculoidei in having a clear globule 142 THE COCCIDIAN PARASITES OF RODENTS in the sporozoites. It differs further from E. kezlint in oocyst size and shape and in having an oocyst polar granule. It differs further from E. musculi in oocyst size and shape, in having an oocyst polar granule, and in sporocyst shape. It differs further from E. schueffneri in oocyst shape and in having an oocyst polar granule. It may differ further from E. hindlet, E. keilini, E. krijgsmanni, E. musculi, and E. schueffneri in other sporocyst characters, but they were so poorly described and illustrated that it is impossible to determine this. It differs further from E. hansonorum in sporocyst shape and size and in the size and shape of the Stieda body. It differs further from E. musculoidei in sporocyst shape, in the character of the Stieda body, and in lacking a sporocyst residuum. This species is named in honor of Dr. Deam H. Ferris, College of Veterinary Medicine, University of Illinois, Urbana. EIMERIA MUSCULOIDEI LEVINE, BRAY, IVENS, AND GUNDERS, 1959 (Plate 29, Fig. 243) Eimeria musculoidei Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts subspherical to ellipsoidal. Oocyst wall pale yel- lowish to yellowish brown, smooth, composed of a single layer about | u thick. Micropyle absent. Thirty-three sporulated oocysts from one mouse measured 17-22 x 15-19 u, with a mean of 19.7 x 16.9 u; their length-width ratios ranged from 1.1-1.3, with a mean of 1.16. One to several oocyst polar granules present. Oocyst residuum absent. Sporocysts lemon-shaped, without Stieda body or with a small, rather flat one. Eight sporocysts measured 10-127 u, with a mean of 10.3x6.9 u; their length-width ratios ranged from 1.4-1.6, with a mean of 1.49. Sporocyst residual granules coarse. Sporozoites lie more or less lengthwise in sporocysts. Sporulation Time: Two to four days at room temperature in Liberia. Schizogony and Gametogony: Schizonts and macrogametes in epithelial cells of upper ileum, usually between host cell nucleus and lumen. Mature schizonts averaged 108 u, with 28 to 36 merozoites. Macro- gametes wtih a central nucleus but without eosinophilic granules. Prepatent Period: Unknown. Type Host: Mus (Leggada) musculoides. Location: Upper ileum. Geographic Distribution: Liberia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in four M. musculoides from Harbel, Liberia. GENUS Eimeria SCHNEIDER, 1875 143 EIMERIA LOPHUROMYSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 * (Plate 29, Fig. 244) Eimeria lophuromysis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts ellipsoidal to ovoid. Oocyst wall yellowish to brownish yellow, smooth, composed of a single layer 0.8 uw thick. Micro- pyle absent. Six oocysts measured 19-23x13-15 u, with a mean of 20.7 x 14.2 u; their length-width ratios ranged from 1.3—1.8, with a mean of 1.45. Oocyst polar granule tiny or absent. Oocyst residuum absent. Sporocysts elongate ellipsoid to slightly ovoid, with very thin wall. Stieda body very small, often no more than a thickening of one end of sporocyst. Sporocysts 116-7 uw, with a length-width ratio of 1.6-1.7. Sporocyst residuum absent. Sporozoites yellowish, with a large, yellowish refractile globule about 3 u in diameter at one end and a small one about 1 u in diameter at the other. Sporozoites lie lengthwise in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lophuromys s. stkapust. Location: Intestinal contents. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA SIKAPUSII LEVINE, BRAY, IVENS, GUNDERS, 1959 (Plate 29, Fig. 245) Eimeria sikapusti Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts subspherical to ellipsoidal. Oocyst wall pale yel- lowish, smooth to slightly pitted, composed of a single layer about | u thick. Oocyst wall often collapses rather quickly in Sheather’s sugar solution. Micropyle absent. Thirteen oocysts from two host animals measured 20-23 x 15-18 u, with a mean of 21.2x17.4 u; their length- width ratios ranged from 1.1—1.3, with a mean of 1.22. One to four large, brownish yellow oocyst polar granules present. Oocyst residuum absent. Sporocysts football-shaped, without Stieda body, 10-12 7-8 u, with a length-width ratio of 1.4—-1.5. Sporocyst residual material abundant, composed of relatively large granules. Sporozoites clear, without refractile globules, lying more or less lengthwise or at the ends of the sporocysts. * The genitive of mys is myis, not mysis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 144 THE COCCIDIAN PARASITES OF RODENTS Schizogony and Gametogony: Unknown, Prepatent Period: Unknown. Type Host: Lophuromys s. sikapust. Location: Intestinal contents. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA LIBERIENSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 (Plate 29 ie.3246) Eimeria liberiensis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts ellipsoidal. Oocyst wall pale yellowish, smooth, composed of a single layer about 0.9 u thick. Micropyle absent. Nine oocysts from two host animals measured 20-27 x 14-20 u, with a mean of 23.7 17.0 u; their length-width ratios ranged from 1.3—-1.7, with a mean of 1.39. Oocyst polar granule present. Oocyst residuum absent. Sporocysts 105 u, with a length-width ratio of 2.0, ellipsoidal, thin-walled, without Stieda body. Sporozoites lie slantwise or curled longitudinally in sporo- cysts, with a compact, granular sporocyst residuum between them. Sporozoites without refractile globules. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lophuromys s. sikapusi. Location: Intestinal contents. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA HARBELENSIS LEVINE, BRAY, IVENS, AND GUNDERS, 1959 (Plate 30, Fig. 248) Eimeria harbelensis Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts ovoid. Oocyst wall smooth, composed of two yellowish brown layers. The inner layer is ellipsoidal, 1.5 u thick at one end, but becoming progressively thinner along the sides until it reaches a thickness of about 0.9 uw at the other end. The outer layer is 1 u thick at the thick end of the inner layer and maintains this width for a little more than two-thirds of the length of the oocyst; it then narrows rapidly, forming a shoulder, and seems to disappear anteriorly. Only a single oocyst was studied; it measured 32x23 u, with a length-width GENUS Eimeria SCHNEIDER, 1875 145 ratio of 1.4. It was not certain whether a micropyle was present; there was a broad, dimplelike indentation 6 u in diameter at the large end of the oocyst which may have been a micropyle. Oocyst polar granule absent. Oocyst residuum absent. Sporocysts 148 u, football-shaped, rather thin-walled, with a medium-sized Stieda body. Sporocyst residuum com- posed of large granules. Sporozoites very granular, with a large, clear globule at one end. Sporozoites lie lengthwise in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lophuromys s. stkapust. Location: Intestinal contents. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA AFRICANA LEVINE, BRAY, IVENS, AND GUNDERS, 1959 (Plate 29, Fig. 247) Eimeria africana Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts subspherical to ellipsoidal. Oocyst wall colorless to pale yellowish, smooth, composed of a single layer about 0.9 wu thick. Oocyst wall collapses rather quickly in Sheather’s sugar solution. Micro- pyle absent. Twenty-five oocysts from four host animals measured 14-21 x 11-16 u, with a mean of 17.0 13.8 u; their length-width ratios ranged from 1.1—1.4, with a mean of 1.23. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, very thin-walled, with small Stieda body. Five sporocysts measured 8-116 u, with a mean of 9.86.0 u; their length-width ratios ranged from 1.5-1.8, with a mean of 1.63. Small amount of finely granular sporocyst residual material usually present. Sporozoites crowded and curled in sporocysts, lying either lengthwise or at the ends of the sporocysts. Sporozoites contain a large, pale yellowish globule at one end and a small one at the other. The oocysts had four sporoblasts when passed. Sporulation Time: Twelve hours at room temperature in Liberia. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lophuromys s. sitkapust. Location: Intestinal contents. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. 146 THE COCCIDIAN PARASITES OF RODENTS EIMERIA KRUIDENIERI LEVINE, BRAY, IVENS, AND GUNDERS, 1959 (Plate 31, Fig. 259) Eimeria kruideniert Levine, Bray, Ivens, and Gunders, 1959: 215-222. Description: Oocysts somewhat ellipsoidal, but narrowed and flattened at both ends. Oocysts sometimes asymmetrical. Oocyst wall yellowish brown, slightly to moderately roughened, sometimes pitted, composed of a single layer about 1.3 u thick at the sides and narrowing to 0.6—0.7 u thick at both ends. Micropyle absent. Twenty-three oocysts from a single host animal measured 27-31 x 19-21 u, with a mean of 28.7 19.4 u; their length-width ratios ranged from 1.3-1.6, with a mean of 1.48. Oocyst polar granule present, often splinterlike. Oocyst residuum absent. Sporocysts 158 u, with a length-width ratio of 1.8, elongate ovoid, rounded at one end and truncate at the other. Stieda body absent. Sporocyst residuum present in some sporocysts but absent in others. Sporozoites somewhat granular, with a large refractile globule at one end. Sporulation Time: Three days at room temperature in Liberia. Schizogony and Gametogony: Macrogametes and oocysts in epithelial cells of jejunum, lying between host cell nucleus and lumen. Macro- gametes contain numerous eosinophilic granules concentrated around their periphery. Prepatent Period: Unknown. Type Host: Lophuromys s. sikapust. Location: Jejunum. Geographic Distribution: Africa (Liberia). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA SCHOUTEDENI VAN DEN BERGHE AND CHARDOME, 1957 (Plate 30, Fig. 249) Eimeria schoutedeni van den Berghe and Chardome, 1957: 1-3. Description: Oocysts ovoid, 14-15 x 11-13 u. Oocyst wall illustrated as a single layer, 0.5 u thick, whitish, presumably smooth. Micropyle absent. Oocyst polar granule presumably absent. Oocyst residuum composed of a small amount of fine powder. Sporocysts 7 x 5.6 u, illustrated without a Stieda body. Sporocyst residuum composed of fine powder. Sporozoites 6x2 u, reniform, with coarse granulations about 0.7 uw in diameter on their surface. Sporulation Time: Five to ten days in 1% chromic acid solution. Schizogony and Gametogony: Unknown. GENUS Eimeria SCHNEIDER, 1875 147 Prepatent Period: Unknown. Sporogony: The sporont divides into two masses, each of which divides again to form two sporocysts. Type Host: Cricetomys dissimilis (hamster rat). Location: Unknown; oocysts found in feces. Geographic Distribution: Africa (Kivu, Congo). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA SP. FANTHAM, 1926 Eimeria sp. Fantham, 1926: 560-570. Description: Oocyst thin-walled and frail, 15-17 x 10-13 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Tatera lobengulae (Lobengula’s gerbille). Location: Weum. Geographic Distribution: South Africa. Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA (?) SP. (YAKIMOFF, GOUSSEFF, AND SUZ’KO, 1945) Eimeria halli (?) Yakimoff. Yakimoff, Gousseff, and Suz’ko, 1945: 172-183. Description: Oocysts ovoid, colorless, without micropyle, 22-31 x 15-23 u, with a mean of 26.3 x 19.9 u; length-width ratio 1.1—-1.6, with a mean of 1.33. The oocysts failed to sporulate. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Nesokia indica huttoni (scaly-toothed rat). Location: Feces. Geographic Distribution: USSR (Tadzhikistan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff, Gousseff, and Suz’ko (1945) found this form in 1 out of 47 N. 7. huttoni from Tadzhikistan. Remarks: On the basis of its size, Yakimoff, Gousseff, and Suz’ko (1945) believed that this form resembled E. nieschulzi (syn., E. halli). However, since it failed to sporulate, it is impossible to assign it a specific name. Host Suborder MYOMORPHA Host Superfamily GLIROIDEA Host Family MUSCARDINIDAE Host Subfamily GLIRINAE 148 THE COCCIDIAN PARASITES OF RODENTS EIMERIA GLIRIS MUSAEV AND VEISOV, 1961 (Plate 39, Fig. 341) Eimeria gliris Musaev and Veisov, 1961a: 1085-88. Description: Oocysts ovoid. Oocyst wall smooth, composed of two layers each 1.25 uw thick, the inner layer dark brown and the outer yellowish. Micropyle absent. Fifty-five sporulated oocysts from three specimens were 14—23 x 12-17 u, with a mean of 21x16 u; their length- width ratios ranged from 1.1—1.5, with a mean of 1.2. Oocyst residuum absent. Oocyst polar granule present. Sporocysts ovoid, 6-10 x 4-8 u, with a mean of 9X7 u. Stieda body absent. Sporocyst residuum composed of many small granules and one large, refractile body. Sporozoites pear- shaped. Sporulation Time: Three days at 25-30C in 2.5% potassium dichro- mate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Glis glis (common dormouse). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Musaev and Veisov (1961a) found this species in three out of eight Glis glis from the Astarin region of Azerbaidzhan. EIMERIA MYOXI GALLI-VALERIO, 1940 Eimeria myoxi Galli-Valerio, 1940: 352-358. Description: Oocysts ovoid, 18x15 u, with a spherical sporont 12 u in diameter. Oocyst wall relatively thin at the micropylar end. Micropyle poorly visible. Sporocysts ovoid, 7.56 u. Sporozoites piriform. Oocyst and sporocyst residua presumably absent, although this is not certain. After discussing the sporocysts, Galli-Valerio (1940) stated simply, “Point de reliquat,” without indicating whether he was referring to the sporo- cyst residuum, the oocyst residuum, or both. Sporulation Time: Seven days on moist filter paper. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Eliomys (syn., Myoxus) quercinus (garden dormouse). Location: Presumably intestine. Geographic Distribution: Europe (Switzerland). Pathogenicity: Unknown. Cross-Transmission Studies: None. GENUS Eimeria SCHNEIER, 1875 149 EIMERIA DYROMIDIS ZOLOTAREYV, 1935 * Eimeria dyromidis Zolotarev, 1935; Musaev and Veisov, 1959a: 535-539. (Plate 30, Fig. 250) Description: Oocysts ovoid, rarely spherical. The ovoid oocysts de- scribed by Zolotarev (1935) measured 16-30 x 13-24 u, with a mean of 22x19 u; the spherical ones were 15-25 u in diameter, with a mean of 19 u. Thirty-three oocysts were measured by Musaev and Veisov (1959a); the ovoid ones were 16—24 x 14-22 u, with a mean of 23x19 u and a mean length-width ratio of 1.2; the spherical ones were 20-22 u in diameter, with a mean of 21 u. Oocyst wall smooth, double-contoured, apparently composed of a single layer 1 u thick; according to Musaev and Veisov (1959a), it is colorless. Micropyle absent. Oocyst residuum absent. Oocyst polar granule absent according to Zolotarev (1935), but present according to Musaev and Veisov (1959a). Sporocysts ovoid; 7— 12 x 6-9 u according to Zolotarev (1935), 10-14. 6-9 u, with a mean of 13 x8 u, according to Musaev and Veisov (1959a). Stieda body apparently absent although Musaev and Veisov (1959a) illustrated the sporocysts as pointed at one end. Sporocyst residuum present. Sporozoites pear- shaped, with a spherical globule at the broad end according to Musaev and Veisov (1959a) but without such a body according to Zolotarev (1935): Sporulation Time: Two days at 25-30C in 2.5% potassium dichromate according to Musaev and Veisov (1959a). Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dryomys nitedula (forest dormouse) (According to Simp- son, 1945, the correct name for this genus is Dryomys ‘Thomas, 1906; however, Thomas (1907) introduced the generic name Dyromys for it under the mistaken impression that Dryomys was preoccupied.) Location: Intestinal contents. Geographic Distribution: USSR (Daghestan and Nakhichevan regions of Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Zolotarev (1935) found this species in all of ten D. nitedula examined from Daghestan. Musaev and Veisov (1959a) found it in one or two of five forest dormice from the Nakhichevan region of Azerbaidzhan. Remarks: Although the oocysts found by Musaev and Veisov (1959a) differed from those described by Zolotarev (1935) from the same host in * The genitive of mys is myis, not midis. Unfortunately, the 1961 International Code of Zoological Nomenclature does not permit a change in spelling of this name, so the error must be perpetuated. 150 THE COCCIDIAN PARASITES OF RODENTS being colorless and in having an oocyst polar granule and a spherical globule in the sporozoites, the former authors considered both to belong to the same species. EIMERIA NACHITSCHEVANICA MUSAEV AND VEISOV, 1959 (Plate 30, Fig. 251) Eimeria nachitschevanica Musaev and Veisov, 1959a: 535-539. Description: Oocysts ovoid. Sixteen sporulated oocysts measured 20— 26 x 17-20 u, with a mean of 24X19 uw and a mean length-width ratio of 1.3. Oocyst wall smooth, colorless, apparently composed of a single layer 1.3 uw thick. Micropyle prominent, with a micropylar cap. Oocyst residuum and polar granule absent. Sporocysts ovoid, with a well- developed Stieda body, 10-13 x 6-9 u, with a mean of 12 <7 u. Sporozoites comma-shaped, with a spherical globule at the broad end. Sporocyst residuum present, consisting of small, scattered granules. Sporulation Time: Three days at 25-30C in 2.5% potassium dichro- mate solution according to Musaev and Veisov (1959a). Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dryomys nitedula (forest dormouse). (See the note under Eimeria dyromidis above for an explanation of this spelling.) Location: Intestinal contents. Geographic Distribution: USSR (mountain region of Nakhichevan Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Host Suborder MYOMORPHA Host Superfamily DIPODOIDEA Host Family DIPODIDAE Host Subfamily DIPODINAE EIMERIA ALLACTAGAE IWANOFF-GOBZEM, 1934 Emend. SVANBAEV, 1956 Eimeria alactagae Iwanoft-Gobzem, 1934: 149-151. Eimeria allactagae: Svanbaev, 1956: 180-191. Description: ‘The oocysts in the type host as described by Iwanoff- Gobzem (1934) are spherical, with a single-layered, thick, uniform wall, measure 22-26 u in diameter, with a mean of 24.4 u, and have an oocyst refractile granule and lack a sporocyst residuum. She did not mention any other morphological characters. ‘The oocysts described by Svanbaev (1956) from Allactaga elator are ovoid, with a double-contoured, smooth, GENUS Eimeria SCHNEIDER, 1875 151 colorless wall 1.7 u thick, measure 22.0 18.4 u, have an oocyst polar granule (which is occasionally absent), oval sporocysts measuring 8.8 x 7.2 u, and lack a micropyle and oocyst and sporocyst residua. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Allactaga major (syn., A. jaculus) (great jerboa). Other Hosts: Allactaga elator (small, five-toed jerboa). Location: Feces. Geographic Distribution: USSR (Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in one out of four A. elator in western Kazakhstan. EIMERIA LAVIERI YAKIMOFF AND GOUSSEFF, 1936 (Plate 40, Fig. 348) Eimeria lavieri Yakimoff and Gousseff, 1936: 447-448; Svanbaev, 1962: 23-39. Description: Oocysts spherical, yellowish orange, 17-18 u in diameter (mean, 17.4 u). Micropyle absent. Sporocysts broadly oval (illustrated as ellipsoidal), 9x 8 uw. Oocyst polar granule and residuum absent. Sporocyst residuum present. Sporozoites comma-shaped, lying lengthwise in sporo- cysts. According to Svanbaev (1962), the oocysts are subspherical or spherical, 19-27 x 18-22 u, with a mean of 21x19 u; their length-width ratio is 1.1—1.2, with a mean of 1.1; the oocyst wall is smooth, 1.0-1.2 u thick, yellow or yellow-brown, without a micropyle; oocyst polar granule is absent; the sporocysts are ellipsoidal or subspherical, 8-9 x 7-8 u, with a mean of 8.57 wu; a sporocyst residuum is present in the form of small granules; and the sporozoites are comma-shaped. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Allactaga major (syn., A. jaculus) (great jerboa). Location: Feces. Geographic Distribution: USSR (Black Azow-Mer, north of Don River; southern Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962) found this coccidium in one out of four A. major in southern Kazakhstan. EIMERIA JOYEUXI YAKIMOFF AND GOUSSEFF, 1936 (Plate 40, Fig. 349) Eimeria joyeuxi Yakimoff and Gousseff, 1936: 447-448; Svanbaev, 1962; 23-39. 152 THE COCCIDIAN PARASITES OF RODENTS Description: Oocysts subspherical, yellow, 24-28 x 21 u (mean 26.4 x 21.4 wu), length-width ratio 1.1-1.3 (mean 1.23). Micropyle illustrated as absent. Sporocysts 13.6 x 10.6 u, broadly ovoid. Oocyst polar granule absent. Oocyst residuum present. Sporocyst residuum absent. Sporozoites comma-shaped, lying lengthwise in sporocysts. According to Svanbaev (1962), the oocysts are ellipsoidal, subspherical, or spherical, 22-29 x 21-27 u, with a mean of 25 x 23 u; their wall is smooth, yellow-brown or orange- brown, 1.3-1.7 u thick, without a micropyle; an oocyst polar granule is absent; the oocyst residuum is in the form of a large, granular mass; the sporocysts are ellipsoidal, subspherical, or spherical, 8-14 7—9 u, with a mean of 108 u; a sporocyst residuum is absent; and the sporozoites are comma-shaped and measure 7-8 x 2—3 u, with a mean of 7X3 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Allactaga major (syn., A. jaculus) (great jerboa). Location: Feces. Geographic Distribution: USSR (Black Azow-Mer, north of Don River; Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1962) found this coccidium in two out of four A. major in southern Kazakhstan. Host Suborder HYSTRICOMORPHA Host Superfamily CAVIOIDEA Host Family CAVITDAE Host Subfamily CAVIINAE EIMERIA CAVIAE SHEATHER, 1924 (Plate 31, Fig. 254; Plate 40, Figs. 350-357; Plate 41, Figs. 358-71) “Kokzidien” of Bugge and Heinke, 1921: 41-42. Eimeria caviae Sheather, 1924: 243-246; Alves de Souza, 1931: 11-14; Henry, 1932: 211-268; Yakimoff, Senyushkina, and Machul’skii, 1940: 391-401; Lapage, 1940: 144-154, 190-202, 242-254, 280-295; RySavy, 1954: 131-174. Description: Oocysts ovoid, ellipsoid, or subspherical, smooth, often brownish, 13-26 x 12-23 u. Micropyle absent. Oocyst wall composed of two layers according to Lapage (1940). Oocyst polar granule absent. Oocyst residuum present according to Lapage (1940), who was the only one to mention this structure specifically. Sporocysts 11-13 x 6-7 wu. Sporo- cyst residuum present. Sporulation Time: Two to three days according to Henry (1932), five to eight days at room temperature according to Sheather (1924) and GENUS Eimeria SCHNEIDER, 1875 153 Bugge and Heinke (1921), or nine to eleven days at 18-22C according to Lapage (1940). Schizogony and Gametogony: These stages have been described in some detail by Lapage (1940) and to a lesser extent by Henry (1932). However, they did not determine the number of merozoite generations. The schizonts produce 12 or less to 32 merozoites which range from 6-16 uw in length. There may or may not be a residual body in the schizont. The young microgametocyte is indistinguishable from a young schizont. When it matures it produces a large number of curved, spindle- shaped microgametes 3 u long, each with two flagella 6-9 u long. The young macrogametes (which Henry and Lapage called macrogametocytes) can be distinguished from schizonts or microgametocytes by the fact that their cytoplasm contains clusters of small spherical granules. These increase in number and size as development proceeds. They stain black with iron hematoxylin, dark blue with Mayer’s haemalum or Delafield’s hematoxylin, or yellow, orange, or red with Heidenhain’s azan or Mal- lory’s triple stain. The red granules flatten out and coalesce to form the outer wall of the oocyst, after which the yellow granules move to the periphery and form the inner wall. Fertilization is believed to take place before the oocyst wall is laid down. Prepatent Period: Eleven to twelve days, according to Henry (1932) and Lapage (1940). Type Host: Cavia porcellus (domestic guinea pig). Other Hosts: Cavia aperea (wild guinea pig, aperea). The only report of E. caviae in wild guinea pigs is that of Alvez de Souza (1931), who found it in a single animal in Brazil. Location: Colon. In addition, Henry (1932) reported that the ceca were occasionally infected and that she found a few isolated coccidia in the small intestines of some guinea pigs. The parasites are found in the epithelial cells of the crypts of Lieberkithn or, in heavy infections, throughout the mucosa. Geographic Distribution: Worldwide. Pathogenicity: Infected guinea pigs most often show no symptoms and are apparently not much affected. In some cases, however, the coccidia may cause diarrhea and even death. In such animals the colon contains small white or yellowish white plaques and petechial hemor- rhages in the mucosa. Sometimes the entire mucosa may be destroyed. Cross-Transmission Studies: None. Prevalence: Bugge and Heinke (1921) found coccidia in 73 per cent of 180 guinea pigs examined in Germany. According to Lapage (1940), E. caviae is common and widely distributed in England. Henry (1932) apparently had no difficulty finding infected guinea pigs in California. 154 THE COCCIDIAN PARASITES OF RODENTS However, Nie (1950) failed to find coccidia in 56 guinea pigs in Penn- sylvania. Host Suborder HYSTRICOMORPHA Host Superfamily CAVIOIDEA Host Family CAVIIDAE Host Subfamily DOLICHOTINAE EIMERIA DOLICHOTIS MORINI, BOERO, AND RODRIGUEZ, 1955 (Plate 30, Figs. 252 and 253) Eimeria dolichotis Morini, Boero, and Rodriguez, 1955: 83-89; Zwart and Strik, 1961: 58-59. Description: Oocysts subspherical to ellipsoidal. Oocyst wall smooth, colorless, illustrated as composed of a single layer. Oocysts 22-26 x 18-19 u, with a mean of 2418.5 u and a mean length-width ratio of 1.3 (16-27 x 14-22 u, with a mean of 21X18 u, according to Zwart and Strik, 1961). Micropyle absent. Oocyst residuum and polar granule absent. Sporocysts approximately 117 u (based on figure), with Stieda body. Sporocyst residuum present. Sporozoites about 9-9.5 u long (8- 135-7 u, with a mean of 11X6 u, according to Zwart and Strik (1961), with a refractile globule. Sporulation Time: Seventy-two to eighty hours. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dolichotis patagonum patagonum (marra). Location: Small intestine, possibly large intestine. Geographic Distribution: Argentina, Netherlands (zoological gardens). Pathogenicity: The marras in which Morini, Boero, and Rodriguez (1955) found this parasite had an enteritis. At autopsy, catarrhal inflam- mation, ulcerated and necrotic areas, and edema of the small intestine were observed. The trichostrongylid, Graphidiodes affinis, was also present. Zwart and Strik (1961) observed a hemorrhagic enteritis of the colon with many oocysts in one of three infected animals. Cross-Transmission Studies: Zwart and Strik (1961) were unable to infect six young, coccidia-free guinea pigs with FE. dolichotis. Prevalence: According to Morini, Boero, and Rodriguez (1955), the marra is almost extinct, being maintained in zoological gardens. Here there is more crowding than normal, so that the transmission rate is high. Host Suborder HYSTRICOMORPHA Host Superfamily CAVIOIDEA GENUS Eimeria SCHNEIDER, 1875 WS 3 Host Family HY DROCHOERIDAE Host Subfamily HYDROCHOERINAE EIMERIA CAPIBARAE CARINI, 1937 (Plate 39, Fig. 342) Eimeria capibarae Carini, 1937: 367-369. Description: Oocysts oval (illustrated as ellipsoidal), 25-33 x 20-28 u, with a mean of 30 26 uw and a length-width ratio of 1.1-1.25. Oocyst wall yellowish, described as double but illustrated as a single layer, 2 u thick, with fine radial striations. Micropyle absent. Oocyst polar granule and residuum absent. Sporocysts ovoid, 14-15 x8 u, with Stieda body and sporocyst residuum. Sporulation Time: Twelve days in 1% chromic acid at room tem- perature. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Hydrochoerus hydrochoerus (syn., H. capibara) (capybara). Location: Feces. Geographic Distribution: Brazil (Sao Paulo). Pathogenicity: Unknown. Cross-Transmission Studies: None. EIMERIA HYDROCHOERI CARINII, 1937 (Plate 39, Fig. 343) Eimeria hydrochoeri Carini, 1937: 367-369. Description: Oocysts ovoid, 20-22 x 16-18 u, with a length-width ratio of 1.1-1.3. Oocyst wall smooth, colorless, 0.8 u thick, described as double- contoured but illustrated as a single layer. Micropyle absent. Oocyst polar granule and residuum absent. Sporocysts ovoid, 10-11 x6-7 u, with a small Stieda body which is not always clearly visible. Sporocyst residuum present, composed of some granules. Sporulation Time: Twelve days in 1% chromic acid at room tempera- ture. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Hydrochoerus hydrochoerus (syn., H. capibara) (capybara). Location: Feces. Geographic Distribution: Brazil (Sao Paulo). Pathogenicity: Unknown. Cross-Transmission Studies: None. Host Suborder HYSTRICOMORPHA 156 THE COCCIDIAN PARASITES OF RODENTS Host Superfamily CAVIOIDEA Host Family DASYPROCTIDAE Host Subfamily CUNICULINAE EIMERIA (?) NOELLERI (RASTEGAIEFF, 1930) BECKER, 1956 (Gen?) noélleri Rastegaieff, 1930: 377-404. (?) Eimeria nolleri: Becker, 1956: 85-139. Description: Oocysts spherical, 19 u in diameter. No sporulated oocysts were seen. Schizogony: Schizonts measuring 10x 7 uw containing merozoites 1.4 u wide were seen in smears of fecal mucus. Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cuniculus (syn., Coelogenus) paca (paca). Location: Feces. Geographic Distribution: South America (zoo in Pyatigorsk, USSR). Pathogenicity: Unknown. Hill (1952) reported that coccidiosis caused the death of a paca (C. paca) in the London zoo, but did not describe the organism. Cross-Transmission Studies: None. Host Suborder HYSTRICOMORPHA Host Superfamily CAVIOIDEA Host Family DASYPROCTIDAE Host Subfamily DASYPROCTINAE EIMERIA PARAENSIS CARINI, 1935 (Plate 39, Fig. 345) Eimeria paraensis Carini, 1935: 342-344. Description: Oocysts spherical or slightly ovoid, 33-40 x 30-35 u (mean given as 36x 36 u in his summary table). Oocyst wall brownish yellow, 2 uw thick, composed of two layers, the outer one with a rough, punctate surface and the inner one radially striated. Micropyle absent in illustra- tion. Oocyst residuum and polar granule absent. Sporocysts ovoid, 2011 u, with a residuum and a slightly prominent Stieda body. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dasyprocta aguti (syn., “Cotia vermelha (Aguti aguti)” of Carini, 1935) (cotia or agouti). Location: Feces. Geographic Distribution: Brazil. Pathogenicity: Unknown. GENUS Eimeria SCHNEIDER, 1875 157 Cross-Transmission Studies: None. Remarks: Carini (1935) gave the host as “Cotia vermelha (Aguti aguti).” However, “Cotia vermelha” is simply the Portuguese for red cotia, and was probably italicized because Carini’s paper was written in French. EIMERIA COTIAE CARINI, 1935 (Plate 39, Fig. 344) Eimeria cotiae Carini, 1935: 342-344. Description: Oocysts ovoid (illustrated as ellipsoidal), 28 or 29x 18 u (there is a discrepancy between Carini’s text description and his summary table). Oocyst wall slightly striated; sometimes quite thick, very slightly rough, and pale brownish yellow; sometimes thinner, smooth, and color- less. Oocyst wall illustrated as composed of a single layer. Micropyle absent. Oocyst residuum and polar granule apparently absent. Sporocysts ovoid, 13x 8-9 u, with Stieda body and residuum composed of scattered granules. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dasyprocta aguti (syn., “Cotia vermelha (Aguti aguti)” of Carini, 1935) (cotia or agouti). Location: Feces. Geographic Distribution: Brazil. Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: See Remarks under E. paraensis for discussion of host name. EIMERIA AGUTI CARINI, 1935 (Plate 39, Fig. 340) Eimeria aguti Carini, 1935: 342-344. Description: Oocysts spherical, 16-17 u in diameter. Oocyst wall thin, smooth, colorless. Micropyle absent in illustration. Oocyst residuum and polar granule apparently absent. Sporocysts 106 u, with small residuum and illustrated with what might be a very small Stieda body. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Dasyprocta aguti (syn., “Cotia vermelha (Aguti aguti)” of Carini, 1935) (cotia or agouti). Location: Feces. Geographic Distribution: Brazil. Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: See Remarks under E. paraensis for discussion of host name. 158 THE COCCIDIAN PARASITES OF RODENTS Host Suborder HYSTRICHOMORPHA Host Superfamily OCTODONTOIDEA Host Family ECHIMYIDAE EIMERIA MYOPOTAMI YAKIMOFF, 1933 (Plate 31, Fig. 255) Eimeria myopotami Yakimoff, 1933: 189-190; Seidel, 1954: 759-764; Hohner, 1955: 337-342; Zajiéek, 1955: 29-38; Seidel, 1956: 117-126; Prasad, 1960a: 207-210. Description: Oocysts ovoid, sometimes ellipsoidal, very seldom spherical or subspherical. Ovoid and ellipsoidal oocysts 22-27 x 18-23 u, with a mean of 24.0—20.4 and a length-width ratio of 1.1-1.25, with a mean of 1.19. Subspherical oocysts 22-25 x 20-23 u, with a length-width ratio of 1.1—-1.2. Spherical oocysts 22-23 u in diameter. The above dimensions are those of Yakimoff (1933). Seidel’s (1954, 1956) measurements differed from these. In his first paper (1954), he gave the mean dimensions as 21.7-24.8 x 18.6 u, and stated that there are two forms; the range of measurements given for the larger form was 21.7-32.5 x 18.6-27.5 u; the range of measurements given for the smaller form was 15.5-18.6 x 12.4— 21.7 u. In his second paper (1956) Seidel gave the same mean dimensions, but combined the two sizes, giving the range as 15.5-32.5 x 12.4-27.5 u. According to Prasad (1960), the oocysts measure 21-26 x 12-17 u, with a mean of 23x14 u. Pellérdy (1960) believed that the ovoid oocysts de- scribed by Yakimoft (1933) were actually those of E. pellucida. Oocyst wall composed of three layers (two, according to Prasad, 1960a), of which the outer one is fragile (relatively thin, according to Prasad, 1960a). Oocyst wall smooth and yellowish brown according to Prasad (1960a). Micropyle absent. (Seidel (1954) stated that there is a micropyle on one side—‘“an einer Langsseite.’’) Oocyst residuum and polar granule absent. Sporocysts pointed at one end, presumably with Stieda body (Prasad, 1960a stated that there was no Stieda body, but his illustration showed one), with sporozoites lying longitudinally and with sporocyst residuum. Sporocysts 9-126 u, according to Seidel (1954). Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Myocastor coypus (nutria, coypu or swamp beaver). Location: Feces. Geographic Distribution: USSR, England, Germany, Czechoslovakia. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Yakimoff (1933) found this species in 20 per cent of 10 nutria on one fur farm, in 59 per cent of 17 nutria on another, and in GENUS Eimeria SCHNEIDER, 1875 159 none of 58 on a third farm, all in Russia. Seidel (1954) and Hohner (1955) saw it in Germany. Prasad (1960a) found it in 2 out of 18 nutrias in England. Zajitek (1955) reported finding it on 4 nutria farms in Czechoslovakia. Remarks: The descriptions of this species by different authors are not entirely in accord. This fact and the lack of certainty of the difference between E. myopotami and E. coypi (see below) make a restudy of the oocysts of these and other nutria coccidia an obvious necessity. EIMERIA PELLUCIDA YAKIMOFF, 1933 Eimeria pellucida Yakimoff, 1933:189-190; Seidel, 1954: 759-764; Seidel, 1956: 117-126; Hohner, 1955: 337-342; Zajitek, 1955: 29-38; Pellérdy, 1960: 327-335. Description: Oocysts ovoid or almost cylindroid, often with a flat micro- pyle. Oocyst wall pale lilac-colored when alive, yellow when dead. Oocyst wall presumably smooth, illustrated with two layers. Living oocysts 30-40 x 20-23 u, with a length-width ratio of 1.6-1.7. Dead oocysts 29-40 x 16-25 u, with a length-width ratio of 1.6-2.0. Seidel (1954, 1956) gave the dimensions as 28-31 x 15.5-19 u. No sporulated oocysts were described by the above authors. Pellérdy redescribed the oocysts of EF. pellucida, but his description differed somewhat from the previous ones. He said that the oocysts are bean- shaped, with one side convex and the other straight or concave; while some of the oocysts appeared oval or cylindrical, their true shape could be demonstrated by rolling them. They measured 21-33 x 12-16 u and had a smooth, colorless, thin wall composed of two layers. A micropyle was present and the oocyst residuum was absent. The sporocysts were piriform and measured 9x6 u. There was a sporocyst residuum. Sporulation Time: Two or three days at room temperature, according to Pellérdy (1960). Schizogony and Gametogony: The endogenous stages have been de- scribed by Pellérdy (1960), but he saw no schizogonic stages. The sexual stages were found in the small intestine from the duodenum through the ileum; the largest numbers were in the ileum. The macrogametes enter the mucous membrane and then come to lie deep under the epithelial layer, where they cause the host cell nuclei to become hypertrophied and flattened. When mature, they measure 22x15 u. Pellérdy was unable to differentiate between the microgametocytes of E. pellucida and those of E. seideli, which were also present. Prepatent Period: Pellérdy (1960) found the prepatent period in an experimentally infected animal which already carried a slight infection was 11 to 12 days. Type Host: Myocastor coypus (nutria). 160 THE COCCIDIAN PARASITES OF RODENTS Geographic Distribution: USSR, Germany, Czechoslovakia, Hungary. Pathogenicity: Pellérdy (1960) stated that E. pellucida may be harmful if it is present in susceptible animals in sufficiently large numbers. Cross-Transmission Studies: None. Prevalence: Yakimoff (1933) found this species in 12 per cent of 17 nutria on a fur farm in Russia; he did not find it on 2 other nutria farms. Seidel (1954) and Hohner (1955) saw it in Germany. Zajicek (1955) reported finding it on 4 nutria farms in Czechoslovakia, and Pellérdy (1960) found it in Hungary. Remarks: Pellérdy (1960) thought that the oval oocysts described by Yakimoff (1933) as those of E. myopotami were actually those of FE. pellucida which were not oriented to show their true shape. The conflicting descriptions of EF. pellucida given above necessitate a restudy of the oocysts of this and other nutria coccidia. EIMERIA COYPI OBITZ AND WADOWSKI, 1937 Eimeria coypi Obitz and Wadowski, 1937: 98-99; Seidel, 1954: 759-764; Seidel, 1956: 117-126. Description: Oocysts ovoid, with a thick, transparent, presumably smooth wall illustrated as composed of a single layer. Oocysts 21-26 x 12— 16 uw, with a mean of 22.8 x 14.7 u, according to Obitz and Wadowski (1937). According to Seidel (1954, 1956), the oocysts are spherical, have a fragile wall, and measure 12.5-17.5 u in diameter. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts el- lipsoidal, 9-12 x 5-7 u, with a mean of 10.66 u, without Stieda body, with sporocyst residuum. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Myocastor coypus (nutria). Location: Feces. Geographic Distribution: Poland, Germany (?). Pathogenicity: According to Obitz and Wadowski (1937), this species seems to be pathogenic in some degree for young animals. Cross-Transmission Studies: None. Prevalence: Obitz and Wadowski (1937) found this species in 40 per cent of 50 nutria examined. Remarks: The description given by Seidel (1954, 1956) for E. coypi differs so much from that of Obitz and Wadowski (1937) that it seems doubtful whether it is the same species. However, Seidel gave no informa- tion other than that presented above, and this is not sufficient to warrant the establishment of a new species. Prasad (1960a) thought that E. coypi was a synonym of £. myopotami, and Pellérdy (1960) suspected that GENUS Eimeria SCHNEIDER, 1875 161 either E. coypi or E. myopotami was a nomen nudum. They may well be right; however, neither saw coccidia which conformed to the original description of E. coypi, so we consider it best to retain this species for the present. EIMERIA SEIDELI PELLERDY, 1957 (Plate 42, Figs. 372-374) Eimeria seideli Pellérdy, 1957: 591; Pellérdy, 1960: 327-335; Pellérdy, 1960a: 389-399. Eimeria (Globidium) fulva Seidel, 1954: 759-764. Eimeria fulva: Seidel, 1954a: 190-191; Seidel, 1956: 117-126. Eimeria (Globidium) perniciosa Sprehn in Seidel, 1954: 759-764; Seidel, 1954a: 190-191. [non] Eimeria fulva Farr, 1963: 336-340. [non] Eimeria (Tyzzeria) perniciosa (Allen, 1936) Seidel, 1954: 759-764. Description: Oocysts spherical, 38-45 u in diameter, or subspherical, 45-48 x 38-42 u, with a length-width ratio of 1.2. Oocyst wall composed of three layers: the outer one (present only in young oocysts) thin and whitish; the middle one light to dark brown, almost opaque, thick, and very rough (“gekérnt”’); and the inner layer colorless and thin. According to Pellérdy (1960), the wall is 2-5 u thick, and the outer layer may peel or chip off easily. Micropyle, oocyst residuum and polar granule absent. Sporocysts piriform to ovoid, 26-2913 u, with a Stieda body. Sporo- zoites bean-shaped, 10 u long according to Seidel (1954). Free sporozoites 14-16 x4 u according to Pellérdy (1960). Sporocyst residuum present. Sporulation Time: Thirty-seven days in 0.5% potassium bichromate solution at room temperature according to Seidel (1954); twelve days according to Pellérdy (1960). Schizogony: Pellérdy (1960a) studied the endogenous development of E. seideli in experimentally infected animals. He found schizogony in a single nutria killed eight days after infection. He thought that there was only a single generation of schizonts. They are found in the epithelial cells of the middle and terminal parts of the small intestine, generally in the depths of the crypts and rarely in the epithelial cells of the villi. They are ovoid to ellipsoidal, measure 15-20 14 u, and contain 10-25 crescentic banana-shaped merozoites which taper to a point at both ends and are up to 16-18 u long. Colonies of schizonts up to 50 u in diameter may occur. Gametogony: Pellérdy (1960a) found the gamonts beneath the epithe- lium throughout the small intestine. The young stages occur beneath the epithelium of the villi, and the older ones are found in the lacteals, within the connective tissue supporting the villi. Occasionally they are deep, near the muscularis mucosae. The host cell nucleus is hypertrophied, 162 THE COCCIDIAN PARASITES OF RODENTS indented by the parasite, first becoming bean-shaped and then rather flattened and elongated. Pellérdy thought that the host cell was an epithelial cell which had been dislodged and had migrated into a sub- | epithelial position after having been invaded, but he was not sure of this and thought it possible that the host cell might be of mesenchymal origin instead. The young macrogametes are 5-6 uw in diameter, with a deep blue nucleus and a red, eccentric karyosome. When they are 15-20 u in diameter, a deep blue, caplike structure appears near the nucleus and seems to migrate toward the periphery of the cell, where its outlines become blurred and it finally disintegrates into granules. Small, deep blue plastic granules appear in the cytoplasm. They enlarge, take on a red stain, migrate toward the periphery of the cell and eventually coalesce to form the oocyst wall. Pellérdy believed that the granules formed from the nuclear cap play no part in oocyst wall formation but simply disappear. According to Seidel (1954, 1956), the mature macro- gametes measure 32-38 x 29-35 u. The young microgametocytes are indistinguishable from the young macrogametes. When they are about 20 uw in diameter, their nucleus divides repeatedly and the resultant nuclei are distributed throughout the cytoplasm. The microgametocytes continue to enlarge, reaching a diam- eter of 50-70 u and containing a very great number of minute nuclei. These then combine in groups to form islets of varying size. The nuclei presumably continue to divide in these islets and come to lie on their periphery. The islets may attain a diameter of 10-15 u, and the micro- gametocytes may eventually reach a diameter of 100 u, although they average 40-60 u and are usually elongated, ovoid, or ellipsoidal. They contain no residuum. The microgametes are dark blue, comma-shaped, and 3-5 wu long. Pellérdy saw no flagella. Prepatent Period: Fourteen days according to Pellérdy (1960a). Type Host: Myocastor coypus (nutria). Location: ‘Throughout small intestine. Geographic Distribution: Europe (Germany, Hungary). Pathogenicity: Seidel (1954, 1956) considered E. seideli to be patho- genic, and Pellérdy (1960) considered it the most pathogenic species in the nutria. Seidel found it in 4- to 5-month-old nutrias which had died with inflammatory changes in the intestine, but other coccidian species, Strongyloides sp. and Trichuris sp. were also present. Pellérdy (1960) produced the disease experimentally in a 6-month-old nutria by feeding about 500,000 oocysts. He described erythema, edema, and a whitish coating on the walls of the duodenum, jejunum, and especially of the GENUS Eimeria SCHNEIDER, 1875 163 ileum, but it is not clear whether these were due exclusively to E. seideli or to it and E. pellucida. Cross-Transmission Studies: None. Prevalence: Seidel (1954, 1956) found this species in 15 of 73 young nutrias which had died on a fur farm in Germany. Remarks: Seidel (1954, 1956) described this species under the name Eimeria (Globidium) fulva. However, this is a homonym of Ezmeria fulva Farr, 1953 from the Canada goose, Branta canadensis, and Pellérdy (1957) assigned the new name E. seideli to Seidel’s species. Seidel (1954, 1956) mentioned that Sprehn in a personal communica- tion had suggested the name Ezmeria (Globidium) perniciosa for this species, but that this name could not be used because it would be a homonym of “Eimeria (Tyzzeria) perniciosa Allen, 1936” of the domestic duck. However, Allen (1936) named her species Tyzzeria perniciosa, and Seidel’s designation appears to be a new combination. This combina- tion, however, is not acceptable. Tyzzeria is not congeneric with Eimeria. Its oocysts contain eight naked sporozoites without any sporocysts, while those of E:meria contain four sporocysts each with two sporozoites. Nevertheless, Seidel’s action makes it impossible to use the specific name, perniciosa, for any species of Eimeria. EIMERIA NUTRIAE PRASAD, 1960 (Plate 31, Figs. 256 and 257) Eimeria nutriae Prasad, 1960a: 207-210. Description: Oocysts broadly ovoid or subspherical. Oocyst wall yel- lowish, pitted like a thimble, composed of a single layer. Micropyle absent. Oocysts 19-23 15-18 u, with a mean of 20+1.9x16+1.4 4u; their length-width ratios ranged from 1.2—1.5, with a mean of 1.3. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, without Stieda body, 10-12 x 4-6 u, with a mean of 11 x5 u. Sporocyst residuum present. Sporozoites spindle-shaped, 9-11 x 2-3 u, with a central nucleus and a refractile globule at one end. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Myocastor coypus (nutria). Location: Feces. Geographic Distribution: England. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Prasad (1960a) found this species in 8 out of 18 fecal sam- ples from nutrias from Norfolk and Sussex, England. 164 THE COCCIDIAN PARASITES OF RODENTS EIMERIA MYOCASTORI PRASAD, 1960 (Plate 31, Fig. 258) Eimeria myocastori Prasad, 1960a: 207-210. Description: Oocyst broadly ovoid. Oocyst wall smooth, colorless, com- posed of two layers of which the outer is thinner than the inner. Micro- pyle conspicuous. Fifty oocysts measured 13-15 x 11-13 u, with a mean of 14x12 u; their length-width ratios averaged 1.1. Oocyst residuum and polar granule absent. Sporocysts cigar-shaped, 9-11 x 3-5 u, with a mean of 10x 4 u. Stieda body absent. Sporocyst residuum present. Sporozoites spindle-shaped, 9-11 x 2.5-3 u, with a refractile globule. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Myocastor coypus (nutria). Location: Feces. Geographic Distribution: England. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Prasad (1960a) found this species in 4 out of 18 fecal samples from nutrias from Norfolk and Sussex, England. Host Suborder HYSTRICOMORPHA Host Superfamily BATHYERGOIDEA Host Family BATHYERGIDAE EIMERIA HETEROCEPHALI N. SP. Eimeria muris Galli-Valerio, 1932 of Porter, 1957: 515-527. [non] Eimeria muris Galli-Valerio, 1932: 129-142. Description: Oocysts ovoid. Small sporocyst residuum present. Sketch of unsporulated oocyst shows no micropyle. No other oocyst character- istics given. Schizogony: Unknown. Gametogony: Macrogametes with “‘chromatoid and _ plastinoid par- ticles.” Prepatent Period: Unknown. Type Host: Heterocephalus glaber (naked mole rat). Location: Epithelial cells of cecum. Double infections of these cells with schizonts and macrogametes were frequent. Geographic Distribution: London Zoo (animal originally from Somali- land or Kenya). Pathogenicity: Unknown. Cross-Transmission Studies: None. Remarks: Porter (1957) assigned this species to Eimeria muris with GENUS Isospora SCHNEIDER, 1881 165 the statement, “Comparison with Eimeria muris from rats, mice, striped rats (Arvicanthus) and other rodents showed such strong correspondence that the parasite of Heterocephalus glaber must be considered to be E. muris, the Naked Mole Rat being a new host.’”’ However, the only valid host of E. muris is Apodemus sylvaticus. Furthermore, even though Porter’s description is extremely sketchy and her drawing crude, they are enough to show that the form from H. glaber is not E. muris, since the latter has a micropyle on a slight projection, while the former does not. No other coccidia have been reported from the genus Heterocephalus (of which H. glaber is the sole species) or from any other member of the superfamily Bathyergoidea, to which it belongs. Porter’s description, sketchy though it is, is enough for the recognition of this form. Hence, we are assigning it the name Eimeria heterocephali. GENUS /JSOSPORA SCHNEIDER, 1881 In this genus, the oocyst has two sporocysts, each containing four sporozoites. ISOSPORA CITELLI LEVINE, IVENS, AND KRUIDENIER, 1957 (Plate 32, Fig. 266) Isospora citelli Levine, Ivens, and Kruidenier, 1957a: 143-144. Description: Oocysts subspherical. ‘Two oocysts measured 22-23 x 21-22 u, with a mean of 22.4 21.5 u and a mean length-width ratio of 1.04. Oocyst wall smooth, composed of two very pale tan layers, the outer one 1 u and the inner one 0.4 u thick. Micropyle absent. Oocyst re- fractile globule present; this looks like an oil droplet; it goes to the top when the oocyst is rolled over. Oocyst residuum absent. Sporocysts broadly lemon-shaped, approximately 15x10 u, with a small Stieda_ body. Sporocysts completely filled by the four sporozoites and residual ma- terial composed of granules of different sizes. The sporozoites lie in no particular order in the sporocysts. In Fig. 266, the sporocysts are shown end-on, so that no Stieda body is visible. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Spermophilus (syn., Citellus) variegatus (rock squirrel). Location: Intestinal contents. 166 THE COCCIDIAN PARASITES OF RODENTS Geographic Distribution: United States (Grand Canyon National Park, Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. ISOSPORA FREUNDI YAKIMOFF AND GOUSSEFF, 1935 (Plate 42, Fig. 375) Isospora freundi Yakimoff and Gousseff, 1935a: 485. Description: Oocysts spherical or subspherical. Spherical oocysts 13- 24 u in diameter, with a mean of 19.8 u. Subspherical oocysts 20-27 x 17-24 u, with a length-width ratio of 1.1—-1.25. Oocyst wall smooth, double- contoured (illustrated as a single layer). Micropyle absent. Oocyst polar granule and residuum absent. Sporocysts in the spherical oocysts 14 x 8-9 UL. Sporocysts illustrated without Stieda body or sporocyst residuum. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Cricetus cricetus (hamster). Location: Feces. Geographic Distribution: USSR (White Russia). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: ‘This species was found in one out of four hamsters. ISOSPORA TERES IWANOFF-GOBZEM, 1934 Isospora teres lwanoft-Gobzem, 1934: 149-151. Description: Oocysts spherical, 24-36 u in diameter. Micropyle absent. Oocyst polar granule present. Oocyst and sporocyst residua absent. Sporo- cysts 16-21 x 8-13 u. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lagurus lagurus. Location: Intestine. Geographic Distribution: USSR (Kazakhstan). Pathogenicity: Unknown. Hyperemia of the intestinal mucosa was ob- served in three rodents infected with this species and J. lagurt. Cross-Transmission Studies: None. Prevalence: Iwanoft-Gobzem (1934) found this species in 3 out of 40 L. lagurus examined, but it is not clear whether she looked for coccidia in animals without intestinal lesions. ISOSPORA LAGURI IWANOFF-GOBZEM, 1934 Isospora laguri Iwanoft-Gobzem, 1934: 149-151; Svanbaev, 1962: 23-39 (?). Isospora laguris [sic] Iwanoff-Gobzem, 1934: Ibid. GENUS Isospora SCHNEIDER, 1881 167 Description: Oocysts ovoid, 24-32 x 16-22 u. Oocyst wall thick. Micro- pyle absent. Sporulated oocysts with large oocyst residuum, without polar granule. Sporocysts 16—21 x 8-13 u. Sporocyst residuum absent. Sporulation Time: Three days. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Lagurus lagurus (steppe lemming). Other Host: Spermophilus (syn., Citellus) maximus (2, see Remarks below). Location: Intestine. Geographic Distribution: USSR (Kazakhstan). Pathogenicity: Unknown. Hyperemia of the intestinal mucosa was ob- served in three rodents infected with this species of J. teres. Prevalence: Iwanoff-Gobzem (1934) found this species in 3 out of 40 L. lagurus examined, but it is not clear whether she looked for coccidia in animals without intestinal lesions. Remarks: Svanbaev (1962) reported finding a coccidium which he called Isospora laguri in 2 (5 per cent) of 43 susliks (Spermophilus maximus) in southern Kazakhstan. The oocysts were ellipsoidal, sub- spherical, or spherical, greenish, yellow-green, or brown, with a smooth wall 1.5-2.0 u thick and without a micropyle. The oocysts measured 18-29 x 18-24 u, with a mean of 23.5 x21 u; their length-width ratio was 1.1. An oocyst polar granule was absent, but an oocyst residuum was present in the form of a large granular ball situated between the sporo- blasts. The sporocysts were ellipsoidal or spherical, 8-13 x 7-11 u, with a mean of 10x 6.4 [sic] u. The sporozoites were comma-shaped, 5—7 x 3-4 u, with a mean of 6x3 u. The above description differs from that of J. laguri from Lagurus lagurus in shape and in having considerably smaller sporocysts. In addi- tion, Spermophilus belongs to a different rodent suborder from that of Lagurus. However, in the absence of more complete descriptions of the forms from both hosts, we are unable to decide whether they belong to the same or different species. Svanbaev (1962) also reported finding a coccidium which he called Isospora laguri in two of seven tamarisk gerbils (Meriones tamariscinus) in southern Kazakhstan. The oocysts were ellipsoidal or subspherical, yellow-green, with a smooth wall 1.5-1.9 u thick and without a micropyle. The oocysts measured 21-30 x 20-26 u, with a mean of 27 x 19.7 [sic] u; their length-width ratio was 1.1 x 1.2, with a mean of 1.35 [sic]. An oocyst polar granule was absent, but an oocyst residuum was present in the form of a granular mass. The sporocysts were ellipsoidal or ovoid, 12-14 7-10 u, with a mean of 139 u. The sporozoites were comma- 168 THE COCCIDIAN PARASITES OF RODENTS shaped or ellipsoidal, 5-6 x 2-3 u, with a mean of 5.5x3 u. A sporocyst residuum was absent. The above description differs from that of J. laguri from Lagurus lagurus in shape and in having smaller sporocysts. In addition, Meriones belongs to a different rodent suborder from that of Lagurus. However, in the absence of more complete descriptions of the forms from both hosts, we are unable to decide whether they belong to the same or different species. ISOSPORA MCDOWELLI SAXE, LEVINE, AND IVENS, 1960 (Plate 31, Fig. 261) Isospora mcdowelli Saxe, Levine, and Ivens, 1960. J. Protozool. 7:61-63. Isospora sp. Saxe, 1952. Proc. Soc. Protozool. 3:13. Description: Oocysts spherical to subspherical. Oocyst wall composed of a single layer. Fifty-three sporulated oocysts measured 9-11 x 8-10 u, with a mean of 109 u; their length-width ratios ranged from 1.0-1.3, with a mean of 1.10. Micropyle absent. Oocyst polar granule and oocyst residuum absent. Sporocysts more or less ellipsoidal; 34 sporocysts measured 6-8 x 4—5 u, with a mean of 7X5 u; their length-width ratios ranged from 1.3-1.7, with a mean of 1.47. Sporocyst residuum present, compact. Sporulation Time: Three to five days at room temperature (24 to 27C) in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Microtus pennsylvanicus (meadow mouse). Location: Cecal contents. Geographic Distribution: United States (Pennsylvania). Pathogenicity: Unknown. Cross-Transmission Studies: None. ISOSPORA BATABATICA MUSAEV AND VEISOV, 1960 (Plate 31, Fig. 262) Isospora batabatica Musaey and Veisov, 1960: 51-61. Description: Oocysts almost spherical or ovoid, colorless. Oocyst wall smooth, double-contoured, composed of a single layer 1 u thick. Micro- pyle absent. Ten sporulated oocysts measured 20-24 x 19-21 u, with a mean of 23x21 u; their length-width ratios ranged from 1.0-1.2, with a mean of 1.1. Oocyst residuum and polar granule absent. Sporocysts ovoid, 9-14 x 6-9 u, with a mean of 138 wu. Stieda body prominent. Sporocyst residuum present, composed of fine granules. Sporozoites ovoid, GENUS Isospora SCHNEIDER, 1881 169 Sporulation Time: Two days in 2.5% potassium dichromate at 25—30C. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Arvicola terrestris (water vole). Location: Large intestine contents. Geographic Distribution: USSR (Azerbaidzhan). Musaev and Veisov (1960) found this species in water voles at Batabat in the alpine zone of the Shakhbuz region of Nakhichevan ASSR, Azerbaidzhan SSR. Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. ISOSPORA ORDUBADICA MUSAEV AND VEISOV, 1960 (Plate 32, Fig. 269) Isospora ordubadica Musaev and Veisov, 1960a: 67-75. Description: Oocysts ovoid or subspherical, colorless. ‘Ten sporulated oocysts measured 18-20 x 14-18 u, with a mean of 2017 u; their length- width ratios ranged from 1.1—1.3, with a mean of 1.2. Oocyst wall smooth, composed of a single layer | u thick. Micropyle absent. Oocyst residuum and polar granule absent. Sporocysts ovoid (illustrated as ellipsoidal), without Stieda body, 10-12 x 8-10 u, with a mean of 12x 10 u. Sporocyst residuum present, in the form of small granules. Sporozoites relatively small, ovoid. Sporulation Time: Two to three days at 25-30C in 2.59% potassium dichromate solution. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Meriones persicus (Persian jird). Location: Intestinal contents. Geographic Distribution: USSR (Nakhichevan region of Azerbaidzhan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Unknown. ISOSPORA URALICAE SVANBAEV, 1956 (Plate 42, Fig. 379) Isospora uralicae Svanbaey, 1956: 180-191. Description: Oocysts ovoid, 26.4 x 22.5 u. Oocyst wall smooth, greenish, 1.6 u thick, described as double-contoured and illustrated as composed of a single layer. Micropyle absent. Oocyst polar granule present. Oocyst residuum absent. Sporocysts ovoid, 13.8 x 9.1 u, illustrated without Stieda body. Sporocyst residuum absent. Sporozoites comma-shaped, 6.9 x 3.7 u. 170 THE COCCIDIAN PARASITES OF RODENTS Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Sporogony: The sporont divides into two masses, each of which be- comes a sporocyst containing four sporozoites. Type Host: Apodemus sylvaticus. Location: Feces. Geographic Distribution: USSR (western Kazakhstan). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Svanbaev (1956) found this species in 4 of 18 A. sylvaticus in western Kazakhstan. ISOSPORA RATTI N. SP. (Plate 32, Fig. 267) Description: Oocysts subspherical. ‘Two sporulated oocysts measured 22-24 x 20-21 yu. Oocyst wall smooth, pale tan to tan, composed of a single layer about 1 u thick. Micropyle absent. Oocyst residuum absent. Two oocyst polar granules present. Sporocysts asymmetrical, broadly ovoid, 16x11 u. Stieda body present. Sporocyst residuum present, com- pact. Sporozoites colorless, not arranged in any particular order in sporocysts. Schizogony and Gametogony: Unknown. No endogenous stages were found in the duodenum, jejunum, anterior ileum, posterior ileum, or cecum of the host animal. Prepatent Period: Unknown. Type Host: Rattus norvegicus (Norway rat). Location: Intestinal contents. Geographic Distribution: North America (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: ‘This species was found in | of I1 wild R. norvegicus in Moultrie County, Illinois. The host animal was trapped on a farm about two miles north of Sullivan, Illinois. Remarks: The species of Jsospora reported from rodents have been described above. None has been reported heretofore from Rattus. I. ratti differs from J. citelli in having a one- rather than a two-layered oocyst wall and in the shape of its sporocysts. It differs from J. frewndi in having an oocyst polar granule, Stieda body, and sporocyst residuum. It differs from J. teres in having a sporocyst residuum. It differs from I. laguri in having an oocyst polar granule and sporocyst residuum and in lacking an oocyst residuum. It differs from 7. mcdowelli in size, in having an oocyst polar granule, and in the shape and size of its sporo- GENUS Wenyonella HOARE, 1933 7A cysts. It differs from J. wralicae in oocyst and sporocyst shape, in sporocyst size, and in having a sporocyst Stieda body and sporocyst residuum. GENUS DORISIELLA RAY, 1930 In this genus, the oocyst has two sporocysts, each containing eight sporozoites. DORISIELLA ARIZONENSIS LEVINE, IVENS, AND KRUIDENIER,, 1955 (Plate 32, Fig. 268) Dorisiella arizonensis Levine, Ivens, and Kruidenier, 1955: 52-53. Description: Oocysts spherical or subspherical. Oocyst wall smooth, composed of two layers, the outer one | u thick and colorless, the inner one 0.5 u thick and pale tan. Micropyle absent. Oocysts 21-28 x 21-22 u, with a mean of 21.8 21.0 u; oocyst length-width ratio 1.00-1.05, with a mean of 1.04. Oocyst residuum absent. One to three oocyst polar granules present. Sporocysts lemon-shaped, thin-walled, 11-13 9-10 u, with a mean of 12.79.2 u. Sporocysts with Stieda body and a few to many scattered, round, clear residual granules or bodies varying in diameter from a fraction of a micron up to 5 u. Sporozoites often oriented more or less lengthwise in sporocysts, sometimes spiralling somewhat around a large, clear central residual body. Sporozoites and residual material completely fill sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Neotoma lepida (desert woodrat). Location: Feces. Geographic Distribution: North America (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. GENUS WENYONELLA HOARE, 1933 In this genus, the oocyst has four sporocysts each containing four sporozoites. 172 THE COCCIDIAN PARASITES OF RODENTS WENYONELLA HOAREI RAY AND DAS GUPTA, 1935 (Plate 42, Figs. 380-382) Wenyonella hoarei Ray and Das Gupta, 1935: 112-113; Ray and Das Gupta, 1937: 117-120. Description: Oocysts spherical, 14—-18.5 u in diameter. Number of layers in oocyst wall not specified. Micropyle absent. Oocyst polar granule and oocyst residuum absent. Sporocysts ovoid, 108 u, with prominent Stieda body. Sporozoites usually lie in pairs around sporocyst residuum with a regular “head to tail” arrangement. Sporulation Time: Seven days in 1% chromic acid at room tempera- ture. Sporogony: The first sign of development occurs after two days, when four lobes project from the center of the sporont. At the tip of each of these are four homogeneous prominences which Ray and Das Gupta (1937) believed to be precociously forming sporozoites. The sporoblasts do not separate until the fourth day. The sporocysts are formed on the fifth day, and become mature on the seventh. Schizogony: Schizogony takes place in the epithelial cells of the small intestine. Six to eight merozoites are formed, but Ray and Das Gupta (1937) were unable to determine whether there was more than one merozoite generation. There are two types of merozoites. One type is 6X2 u, hyaline, with a granule near the nucleus which stains darkly with hematoxylin. The other type is 8x2 u, more opaque, and with numerous, irregularly scattered, darkly staining granules. Both types are broad and rounded at one end and pointed at the other; the pointed region stains darkly with hematoxylin. Ray and Das Gupta (1937) believed that the smaller merozoites gave rise to microgametocytes and the larger ones to “macrogametocytes.”’ Gametogony: The young microgametocytes are 6—8 u in diameter. After they have grown, a large number of nuclei are found at their periphery. Each nucleus lies in a cytoplasmic protuberance and has a basal granule above it. Two flagella are formed; in the mature microgamete one of these is directed backwards. A large number of microgametes is formed, leaving a small cytoplasmic residuum. The microgametes break from the residuum and swim actively, congregating at the “micropylar” end of the macrogametes. Ray and Das Gupta (1937) did not describe macrogamete formation in detail. They stated merely that as the “macrogametocyte” increased in size, the number of granules in the cytoplasm also increased, the nucleus shifted toward one pole, and its karyosome enlarged and then disintegrated. GENUS Wenyonella HOARE, 1933 173 Prepatent Period: Unknown. Type Host: Sciurus sp. (Indian squirrel). Location: Small intestine. Geographic Distribution: India (Bengal). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was seen in three squirrels. WENYONELLA UELENSIS VAN DEN BERGHE, 1938 (Blater42 ies 5717)) Wenyonella uelensis van den Berghe, 1938: 275-277. Description: Oocysts described as ovoid but illustrated as ellipsoidal, 26-30 x 19-20 u. Oocyst wall thick. Micropyle absent. Oocyst polar gran- ule absent. Oocyst residuum present at first, but disappears almost com- pletely by fifth day of sporulation. Sporocysts 11 x8 u, illustrated as ellipsoidal, without Stieda body. Sporocyst residuum present, insignifi- cant. Sporozoites banana-shaped. Sporulation Time: Five days (?) in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Funisciurus anerythrus. Location: Probably intestine. Geographic Distribution: Africa (Belgian Congo). Pathogenicity: Unknown. Cross-Transmission Studies: None. WENYONELLA PARVA VAN DEN BERGHE, 1938 (Plate 42, Fig. 378) Wenyonella parva van den Berghe, 1938: 275-277. Description: Oocysts subspherical, 15x13 u. Oocyst wall thick, com- posed of two layers. Micropyle absent. Oocyst polar granule absent. Oocyst residuum absent. Sporocysts 8x5 u, illustrated as ellipsoidal, without Stieda body. Sporocyst residuum absent. Sporulation Time: Seven days in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Tamiscus emini. Location: Probably intestine. Geographic Distribution: Africa (Belgian Congo). Pathogenicity: Unknown. Cross-Transmission Studies: None. 174 THE COCCIDIAN PARASITES OF RODENTS GENUS CARYOSPORA LEGER, 1904 In this genus the oocyst has a single sporocyst containing eight sporozoites. CARYOSPORA MICROTI SAXE, LEVINE, AND IVENS, 1960 (Plate 31, Fig. 263; Plate 32, Fig. 264) Caryospora microti Saxe, Levine, and Ivens, 1960: 61-63. Caryospora sp. Saxe, 1952: 13. Description: Oocysts spherical to subspherical. Oocyst wall composed of a single layer. Micropyle absent. Twelve sporulated oocysts measured 9-10.5 x 8.5-10 u, with a mean of 109 u; their length-width ratios ranged from 1.0-1.2, with a mean of 1.08. Oocyst polar granule and oocyst residuum absent. Sporocysts spherical to subspherical. Twelve sporulated sporocysts measured 7—8.5 x 6—7 u, with a mean of 76.5 u; their length-width ratios ranged from 1.0-1.4, with a mean of 1.12. Stieda body absent. Sporocyst residuum present, compact. Sporulation Time: Ten days or more at room temperature (24 to 27C) in 1% chromic acid. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Microtus pennsylvanicus (meadow mouse). Location: Cecal contents. Geographic Distribution: United States (Pennsylvania). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: This species was found in a single meadow mouse. GENUS TYZZERIA ALLEN, 1936 In this genus the oocyst contains eight naked sporozoites. TYZZERIA PEROMYSCI LEVINE AND IVENS, 1960 (Plate 32, Fig. 265) Tyzzeria peromysci Levine and Ivens, 1960: 207-212. Description: This species was reported from two species of Peromyscus. Since there are minor differences in the appearance of the oocysts from these hosts, each form is described separately. GENUS Tyzzeria ALLEN, 1936 175 Form from type host, Peromyscus maniculatus: Oocysts ellipsoidal. Oocyst wall colorless to very pale yellowish, smooth, composed of a single layer about 0.6 « thick. Micropyle absent. Forty sporulated oocysts from two host animals measured 11—14 x 9-11 u, with a mean of 12.6 x 10.1 u; their length-width ratios ranged from 1.1—-1.4, with a mean of 1.24. One or two polar granules present. Sporozoites banana-shaped, crescent- shaped, or lanceolate, about 3.5 u wide and 9-11 u long. Sporozoites clustered together in a ball without orientation in any particular direc- tion, sometimes with | or 2 sporozoites separated from the others. Sporo- zoites often with fine granules in cytoplasm except in central region where the nucleus is; sporozoites sometimes without granular cytoplasm. No oocyst residuum seen, but it is possible that a small one might have been concealed inside the ball of sporozoites. Form from P. leucopus: Oocysts ellipsoidal. Oocyst wall very pale yellowish, smooth, composed of a single layer about 0.6 uw thick. Micro- pyle absent. Twenty sporulated oocysts measured 14-17 x 11-12 u, with a mean of 15.2x11.2 u; their length-width ratios ranged from 1.3-1.5, with a mean of 1.36. One or two polar granules present. Sporozoites banana-shaped, crescentic, or lanceolate, clustered together in a ball without orientation in any particular direction, sometimes with | or 2 sporozoites separated from the others. A small, compact oocyst residuum 3—4 u in diameter was seen in some oocysts in which the sporozoites were sufficiently well separated, and one may have been present but hidden by the sporozoites clustered around it in most or all of the other oocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus maniculatus (deer mouse). Other Host: Peromyscus leucopus (deer mouse). Location: Intestinal contents. Geographic Distribution: United States (Illinois). Pathogenicity: Unknown. Cross-Transmission Studies: None. Prevalence: Levine and Ivens (1960) found this species in two of nine P. maniculatus and one of seven P. lewcopus from the vicinity of Sullivan, Illinois. Remarks: The form from P. maniculatus differs from that from P. leucopus only in that it is slightly smaller and broader and in that no oocyst residuum was seen in it. However, as mentioned above, an oocyst residuum might have been present in the center of the ball of sporozoites. These differences do not appear sufficient to justify the establishment of separate species, although future research may reveal differences in the endogenous stages or host-parasite relations which would do so. 176 THE COCCIDIAN PARASITES OF RODENTS GENUS CRYPTOSPORIDIUM TYZZER, 1907 In this genus the oocyst contains four naked sporozoites. CRYPTOSPORIDIUM MURIS TYZZER, 1907 (Plate 43, Figs. 383-403) Cryptosporidium muris Tyzzer, 1907: 12-13; Tyzzer, 1910: 487-510. Description: Oocysts ovoid, ellipsoidal, or spherical, with a knoblike attachment organ at some point on their surface. The attachment organ is bluntly conoidal and often has a delicate, threadlike process extending outward from its apex. Oocyst wall smooth, apparently composed of a single layer. Oocysts measure approximately 7x5 u. Sporocysts absent. Micropyle absent. Oocyst residuum present. Schizogony: The schizonts, like the oocysts and macrogametes, have an attachment organ. Their cytoplasm stains dark blue with Wright's or Giemsa’s stain, and contains no granules. Their maximum size is 7X 6 u. The nucleus divides three times to form eight daughter nuclei. These become grouped at the end of the schizont farthest from its point of attachment. The cytoplasm then cleaves, and eight banana-shaped mero- zoites are formed. They extend from one pole of the schizont to the other, and have a slight spiral twist. A globule of residual material is left near the organ of attachment. The merozoites are 5-8 u thick, with an oval nucleus about one-third of the way from the large end and often a tiny hematoxylin-staining granule between it and the small end. Some merozoites are short and thick, while others are relatively slender. The merozoites break out of the schizont, become attached to the sur- face of the gastric epithelium by one end and round up. A fat globule appears in the merozoite after it becomes attached to the epithelium. The number of asexual generations is unknown. Gametogony: The microgametocytes are similar in morphology and mode of development to the schizonts, but are smaller and contain a relatively large amount of chromatin. They have a thin limiting membrane, an attachment organ, and one or more fat globules. Their maximum size is 5 x 3.5 u. The nucleus divides repeatedly to form 16 chromatin masses which become more and more dense, elongate, and push out from the surface of the microgametocyte. A relatively large residual mass is left. The microgametes are composed of a rod of chromatin with a small amount of faintly staining material about as long as the chromatin rod trailing off from it; there are no flagella. The macrogametes can be distinguished from the schizonts and micro- gametocytes by the presence of oval refractile granules which stain GENUS Cryptosporidium TYZZER, 1907 17 ~w reddish brown to deep purple with iodine and red with Giemsa’s stain after formol fixation. The macrogametes are spherical to ellipsoidal, with an attachment organ at one end which makes them appear flask-shaped. Their limiting membrane becomes thick and impervious to blood stains when they are mature. Fertilization was not observed, although micro- gametes were occasionally found on the surface of the macrogametes. After fertilization the limiting membrane becomes thicker and forms a dense cyst wall. The nuclear changes could not be followed. Sporogony: The protoplasm of the oocyst becomes massed at the distal end of the organism, while the remaining portion is occupied by food granules. The protoplasm segments into four masses which elongate and develop into sporozoites, leaving a mass of residual material containing iodophilic granules, lipoid globules, and some material which stains deeply with Giemsa and hematoxylin. Sporogony takes place within the host. Free sporozoites are often found in the stomach. They are boomer- ang-shaped, with a thin, pointed anterior end with a rod-shaped nucleus near it. They can be distinguished from merozoites by their greater length and by the shape of their nuclei. Type Host: Mus musculus (house mouse) (reported only from labora- tory strains and not from wild mice). Other Hosts: Elton et al. (1931) reported finding C. muris in Apodemus sylvaticus and Clethrionomys glareolus in England, but gave no descrip- tions. Location: Stomach. C. muris develops either attached to the surface of the epithelium of the gastric glands or free in their lumen. It never penetrates the host cells, but is attached to them by means of the attach- ment organ described above. Geographic Distribution: United States. Tyzzer (1910) believed this parasite to have a wide geographic distribution, but it has apparently not been reported from house mice elsewhere, although forms seen by Clarke (1895) and Wenyon (1926) in England may have been this species. In addition, Elton et al. (1931) reported finding C. muris in Apodemus sylvaticus and Clethrionomys glareolus in England but did not describe it. Pathogenicity: According to Tyzzer (1910), C. muris is only slightly pathogenic. The only marked histological change he found was dilata- tion of the gastric glands in severe infections and a slight increase in the collections of lymphoid cells, but he saw nothing in the nature of acute inflammation. Cross-Transmission Studies: Vyzzer (1910) infected laboratory mice experimentally with C. muris, but was unable to infect a white Rattus norvegicus with it. 178 THE COCCIDIAN PARASITES OF RODENTS Prevalence: According to Tyzzer (1910), this parasite is frequently found in laboratory mice. CRYPTOSPORIDIUM PARVUM ‘TYZZER, 1912 (Plate 43, Figs. 404-406; Plate 44, Figs. 407-422) Cryptosporidium parvum Tyzzer, 1912: 394-412. Description: Oocysts ovoid or spherical, 4—5 x 3 u. Oocyst wall smooth, composed of a single layer, with a small, knoblike attachment organ which may be blunt and rounded, may have a blunt conoidal point, or may sometimes be drawn out and slender. A delicate shred or filament which may possibly be derived from the host cell is occasionally found projecting from the attachment organ. Micropyle absent. Sporocysts absent. Oocyst residuum present. Sporozoites slender, bow- or boomer- ang-shaped, 5.5-6 uw long, with rod-shaped, slender nucleus near the anterior end. Schizogony: Schizonts at first no more than 1.5 u in diameter, with an attachment organ. They are attached to the cell surface or embedded in its striated border. They frequently contain one or two small vacuoles. The mature schizonts are 3-5 u in diameter. The nucleus divides three times to form eight daughter nuclei, which come to lie at the surface of the organism opposite the attachment organ. Cytoplasmic cleavage then takes place, and eight falciform merozoites and a small residual mass are formed. The merozoites are 2.5-5 uw long and 0.5-0.7 u wide, with a spherical to ellipsoidal nucleus near the thicker end and often with a granule between the nucleus and the posterior end. There is a minute vacuole in the cytoplasm of the larger merozoites. Gametogony: The microgametocytes are smaller than the schizonts but contain a relatively large amount of chromatin. They have an attach- ment organ. They give rise to 16 tiny microgametes and a spherical mass of residual material which frequently contains a vacuole. The microgametes are composed of chromatin rods about | u long and not more than 0.4 u wide and some faintly staining material with no distinct form. No flagella were seen. The macrogametes are larger than the schizonts and microgametocytes and can be distinguished from them by the presence of tiny, refractile granules which stain reddish brown with iodine and darkly with Giemsa’s stain. They have a thin, dense limiting membrane and an attachment organ. Tyzzer (1912) saw a rod-shaped mass of chromatin in addition to the nucleus in a few small macrogametes, which he interpreted as a micro- gamete nucleus following fertilization. Sporogony takes place inside the host. GENUS Klossia SCHNEIDER, 1875 179 Type Host: Mus musculus (house mouse) (reported only from labora- tory strains and not from wild mice). Location: Small intestine from the pylorus to the ileocecal valve; usually more numerous below the region of Briinner’s glands. The para- sites are usually distributed over the entire surface of the intestinal villi, but are never found in the glands. They are less frequent near the bases of the villi than near the tips, where they often occur in large numbers in the irregular depressions which indent this part of the epithelial surface. All growing forms are attached to the surface of the mucosa, where they either indent or are buried in the striated border of the cells. They do not penetrate the cell cytoplasm. Tyzzer (1912) considered C. parvum to form a link between the strictly extracellular C. muris and the other coccidia which live within the host cell cytoplasm. Geographic Distribution: United States. Pathogenicity: According to Tyzzer (1912), the injury due to this species is insignificant, probably amounting to no more than a loss of enough material from the striated border of the cell to accommodate the organ- isms. There is no inflammation. Cross-Transmission Studies: None. Prevalence: According to Tyzzer (1912), this parasite is frequently found in laboratory mice. GENUS KLOSSIA SCHNEIDER, 1875 In this genus the oocysts contain many spherical sporocysts, each with three to ten sporozoites. The macrogametes are not vermiform. KLOSSIA PERPLEXENS LEVINE, IVENS, AND KRUIDENIER, 1955 (Plate 31, Fig. 260) Klossia perplexens Levine, Ivens, and Kruidenier, 1955a: 623-629. Description: Oocysts ellipsoidal, with smooth wall composed of two layers. Outer layer colorless, transparent, about 2 u thick. Inner wall pale brown, 0.5 uw thick. Micropyle absent. Oocysts 42-53 «x 35-44 u, with a mean of 47.9 x 39.9 u; standard deviation of length, 3.53, of width 2.99. Oocyst length-width ratio ranges from 1.17—1.22, with a mean of 1.20 and a standard deviation of 0.018. Oocyst polar granule absent. Oocyst residuum composed of about a dozen round, colorless, homogeneous 180 THE COCCIDIAN PARASITES OF RODENTS globules 3-6 uw in diameter scattered among the sporocysts. Twelve sporocysts present in each of four oocysts carefully counted. Sporocysts packed so tightly in oocysts that counting them is difficult. Sporocysts spherical, without Stieda body, with colorless wall 0.5 u thick. Sporo- cysts 13-14 u in diameter, with a mean of 13.3 u. Each sporocyst contains four sausage-shaped, colorless sporozoites arranged in no_ particular order. Colorless, spherical granules about 1.2—1.5 uw in diameter in con- stant Brownian movement fill the remaining space in the sporocysts. One or two larger granules about 3 u in diameter similar to those in oocysts are sometimes present in sporocysts. Schizogony and Gametogony: Unknown. Prepatent Period: Unknown. Type Host: Peromyscus maniculatus (deer mouse). Location: Feces. Geographic Distribution: North America (Arizona). Pathogenicity: Unknown. Cross-Transmission Studies: None. GENUS KLOSSIELLA SMITH AND JOHNSON, 1902 In this genus typical oocysts are not formed, but a number of sporocysts, each with many sporozoites, develops within a membrane which is per- haps laid down by the host cell. KLOSSIELLA MURIS SMITH AND JOHNSON, 1902 (Plate 45, Figs. 423-429; Plate 46, Figs. 430-437) Klossiella muris Smith and Johnson, 1902: 303-316; Woodcock, 1904: 153-163; Brugnatelli, 1908: 121-126; Sangiorgi, 1911: 523-526; Stevenson, 1915: 127-135; Maisin, 1923: 1219-21; Twort and Twort, 1923: 219-242; Bonne, 1925: 1190-92; Wenyon, 1926; Sternberg, 1929: 419-421; Cannarella, 1931: 670-684; Gard, 1945: 427-434: Otto, 1957: 41-48. Klossiella sp. Pfeiffer, 1891. Coccidium klossiella Elaut, 1932: 1012-14. Coccidia. Smith, 1889: 211-217. Description: It is not certain whether true oocysts are present in this genus. The sporocysts develop from a sporont within a vacuole in the host cell. They are surrounded by a membrane, but this may be a host cell structure. Wenyon (1926) stated, “It is doubtful if a true odcyst is formed.” However, Reichenow (1953) considered oocysts to be present. GENUS Klossiella SMITH AND JOHNSON, 1902 181 The morphological features of the various stages in the life cycle are given below. Schizogony: Schizogony takes place in the glomeruli of the kidneys. Smith and Johnson (1902) described schizogonic forms in the epithelium —usually the visceral layer—of Bowman’s capsule. Stevenson (1915) found two types, one in the endothelial cells of the kidney arterioles and the other in the cells forming the capillaries in the kidney glomeruli and also in the endothelial cells of the kidney arterioles. Otto (1957) found schizogonic forms almost exclusively in cysts in the glomeruli; they were primarily in the endothelial cells. Bonne (1925) reported them in addi- tion in the endothelial cells of the capillaries of the lungs and spleen. Twort and Twort (1923) made microscopic examinations of the kidneys of several hundred mice, the adenomatous and hyaline thyroid glands of 150 mice, and other organs of a large number of mice in the course of a study of diseases of 60,000 experimental mice in England. They found K. muris in 90 per cent of the kidneys, 7 per cent of the brains, 15 per cent of the suprarenal glands, 5 per cent of the lungs, 15 per cent of the thyroids, 2 per cent of the spleens and 1 per cent of the lymph glands and pituitary glands. The protozoa were usually in the capillaries. Infection presumably takes place by ingestion of sporocysts, and the sporozoites presumably pass by way of the blood stream. Stevenson (1915) found a small gregariniform body about 12 u long in the peripheral blood of a mouse which had no parasites in the kidneys but which had lived in a cage with others that were passing sporocysts. Of the two types of schizogony described by Stevenson (1916), the type which he considered to be the earlier occurs in the endothelial cells of the kidney arterioles. In this type each schizont forms 8 to 12 mero- zoites. The second type of schizogony is much more common, and occurs in the endothelial cells of the kidney capillaries and arterioles; this is also the type which Smith and Johnson (1902) found in the epithelium of Bowman’s capsule and Bonne (1925) reported from the lungs and spleen. In this type 40 to 60 more or less falciform merozoites each approximately 7x2 u are formed. The mature schizont occupies quite a large portion of Bowman’s capsule. It breaks, releasing the merozoites into the lumen of the kidney tubule. Gametogony: The merozoites enter the epithelial cells of the con- voluted tubules of the kidney where they become gamonts and where gametogony and sporogony take place. Gametogony has been described by Smith and Johnson (1902), Stevenson (1915), Sternberg (1929), and Gard (1945), to whom reference may be made for other reports. A macrogamete and microgametocyte are found together within a vacuole in the host cell. The microgametocyte divides to form two microgametes. 182 THE COCCIDIAN PARASITES OF RODENTS Fertilization then takes place, and the fertilized macrogamete or zygote becomes a sporont or “mother sporoblast’” which may reach a diameter of 40 u. Sporogony: The sporont contains 10 to 20 plastin granules about 1.5 u in diameter. It divides by multiple fission to form 12 to 16 sporoblasts and a residual body. Within each sporoblast about 25 to 34 banana- shaped sporozoites and a sporocyst residuum are formed by multiple fission. The resulting sporocyst is subspherical to spherical, approxi- mately 16x13 u, and has a very thin wall. The sporozoites lie side by side in the sporocyst. As mentioned above, all authorities do not agree as to whether there is a true oocyst. The sporocysts are released into the lumen of the kidney tubules by rupture of the host cell, and pass out in the urine. Prepatent Period: Unknown. It is presumably long, since sporocysts are not shed by mice less than six months old. Type Host: Mus musculus (house mouse). Location: Schizogony in epithelial cells of capillaries and arterioles in kidney, lungs, spleen, and other organs; gametogony and sporogony in epithelial cells of convoluted tubules of kidney. Geographic Distribution: Probably worldwide. This species has been reported from the United States, England, France, Germany, Belgium, Holland, Italy, Austria, Roumania, and Sweden. Pathogenicity: According to Smith and Johnson (1902), in heavy in- fections the kidneys have minute greyish spots over their entire surface. These spots represent foci of necrobiotic changes in the cortex, followed by marked cell proliferation. Otto (1957) described a perivascular, fol- licular, lymphocytic infiltration in the region of the medullary cortex which he considered of diagnostic significance. The epithelium of the infected kidney tubules is destroyed, and the tubules come to look like elongated, contorted bags filled with sporocysts. In the later stages, inter- tubular foci are formed by round cell proliferation. There is no inflam- matory reaction. Giant cells and polymorphonuclear leucocytes are ab- sent. Fatal infections have not been reported. Cross-Transmission Studies: Cannarella (1931) claimed to have trans- mitted K. muris to the guinea pig. She fed sporocysts in mouse urine to a single guinea pig, and found Klossiella in its kidneys 30 days later. However, she used no controls, and this claim has not been confirmed. Prevalence: The true prevalence of K. muris in laboratory mice is unknown. In the laboratory colonies in which it was found, from 20-100 per cent of the mice were infected. Elaut (1932) studied the relation of age to incidence of infection. He found no infections in the kidneys of mice less than 6 months old, but found that the incidence increased GENUS Klossiella SMITH AND JOHNSON, 1902 183 with age up to 100 per cent in animals one year old or more. ‘Twort and Twort (1923) stated that they expected to find evidence of Klossiella nephritis in at least 90 per cent of their animals by the time they were 12 months old, and that as they grew older, probably very few animals escaped infection. Banciu, Dincolesco, and Petrovici (1958) found it in the kidneys of 3 per cent of 397 white mice bred in their laboratory in Roumania. The only report of K. muris in wild house mice was that of Smith and Johnson (1902), who found it in a considerable number caught in their animal room. KLOSSIELLA COBAYAE SEIDELIN, 1914 Klossiella cobayae Seidelin, 1914: 553-564; Stevenson in Wenyon, 1926. Klossiella sp. Pearce, 1916: 431-442; Bonciu et al., 1957: 131-143. Klossia caviae Sangiorgi, 1916: 49-53. Coccidium. Pianese, 1901: 350-367. Coccidia. Alves de Souza, 1931: 11-14. Description: The following description of the various stages in the life cycle is based primarily on that of Stevenson as given in Wenyon (1926). Schizogony: The sporozoites are released from the sporocysts in the lumen of the guinea pig intestine and make their way to the endothelial cells of the capillaries, where the first schizogonic multiplication takes place. The capillaries most commonly affected are those of the kidney glomeruli, but those of other organs may also be involved. Each sporo- zoite forms a small rounded body within the endothelial cell. This schizont attains a diameter of about 2.5 u. Its nucleus then divides re- peatedly until eight are present. Merozoites are formed around the nuclei, where they lie in a vacuole and make the host cell bulge into the lumen of the capillary. The merozoites measure 21 u. They are released into the blood by rupture of the host cell and invade other endothelial cells where the cycle is presumably repeated. Eventually some of them enter the lumen of the kidney tubules and pass to the convoluted portion, where they penetrate the tubule cells. They become schizonts which produce about a hundred merozoites. The host cell is greatly enlarged, almost filling the lumen of the tubule. It ruptures and the merozoites are released into the lumen. Gametogony: The merozoites pass down to the straight kidney tubules or loops of Henle, where they enter the endothelial cells. They occur here typically in pairs, one being a microgametocyte and the other a macrogamete. Sometimes there are two or even three microgametocytes associated with a macrogamete. The microgametocyte divides once to 184 THE COCCIDIAN PARASITES OF RODENTS form two microgametes, one of which then fertilizes the macrogamete. The resultant zygote grows, becoming a sporont and causing its host cell to swell until the lumen of the kidney tubule is completely occluded. At this time the sporont has a diameter of 30-40 u. Sporogony: The sporont nucleus divides repeatedly during the sporont’s growth to form 30 or more daughter nuclei. A sporoblast is formed around each one by a process of budding from the sporont surface. A residuum is left. According to Stevenson and Wenyon, it is doubtful if a true oocyst is formed, but Reichenow (1953) considered that there is one. The sporoblasts become slightly elongated, and a sporocyst wall is formed around each. About 30 sporozoites are formed in each sporocyst. The sporulated sporocysts rupture the host cell, pass down the kidney tubules and escape in the urine, becoming available to infect another host. Prepatent Period: Unknown. Type Host: Cavia porcellus (domestic guinea pig). Other Host: Cavia aperea (wild guinea pig, aperea). The only report of this species in wild guinea pigs is that of Alves de Souza (1931), who found it in a single animal in Brazil. Location: The first schizogony takes place in the endothelial cells of capillaries in the kidney and other organs. The last schizogonic cycle takes place in the endothelial cells of the convoluted portion of the kidney tubules. Gametogony and sporogony take place in the endothelial cells of the straight kidney tubules and loops of Henle. Geographic Distribution: Presumably worldwide. Seidelin (1914) found this species in two guinea pigs in Nigeria, Pianese (1901) and Sangiorgi (1916) found it in Italy, Stevenson in Wenyon (1926) in England, Bonciu et al. (1957) in Roumania, Alves de Souza (1931) in a wild guinea pig in Brazil, Pearce (1916) in 12 guinea pigs from Pennsylvania and New Jersey, and C. C. Morrill and the authors (unpublished) found it in laboratory guinea pigs in Ilinois. Pathogenicity: K. cobayae apparently causes no clinical signs. Accord- ing to Pearce (1916), the pathologic changes in the kidney are slight. They consist of an irregular accumulation of round cells and fibroblasts around some of the glomeruli. There is slight involvement of the labyrinthine tissue adjacent to the glomeruli and apparently none at all in the lower or inner half of the kidney cortex where most of the parasites are found. On the other hand, Bonciu et al. (1957) said that 88 per cent of their cases had slowly developing subacute nephritis with degenera- tive lesions, either simple or combined with proliferative lesions in- filtrated with mononuclear cells, fibroblasts, and sometimes plasmocytes DISCUSSION 185 and eosinophiles; sclerotic chronic nephritis was present in 2 per cent. They also described nodular perivascular mononuclear infiltration of the lungs and deposition of blood pigment in the red pulp of the spleen and in the reticular zone of the adrenal cortex. They found parasites only in the kidneys, however. Cross-Transmission Studies: None. Prevalence: In a study of 976 guinea pigs in Roumania, Bonciu et al. (1957) found Klossiella in an average of 60.5 per cent of the adult animals (its prevalence ranging from 13 to 65 per cent in different groups), and in 17 per cent of the aged guinea pigs. The youngest infected animals were three weeks old. The maximum infection rate (91 per cent) occurred in the summer and the minimum (43-49 per cent) in the winter. KLOSSIELLA SP. (HARTMANN AND SCHILLING, 1917) NOLLER, 1921 Hepatozoon jaculi (2) Hartmann and Schilling, 1917. Klossiella sp. N6ller in Reichenow, 1921; Wenyon, 1926. Description: Uncertain. Type Host: Jaculus jaculus (jerboa). Remarks: Hartmann and Schilling (1917) described schizogonic forms in the endothelial cells of the capillaries of the lung, liver, spleen, and other internal organs of the jerboa which they believed to be part of the life cycle of Hepatozoon balfouri (syn., H. jaculi). However, Noller examined their preparations and informed Reichenow (1921) that they were developmental stages of a Klossiella. Wenyon (1926) also saw this parasite in the jerboa from the Sudan in 1906 and agreed that it is not H. balfourit but a Klossiella. The only forms he saw appeared to be schizogonic stages in the kidney tubules. DISCUSSION In ‘Tables | through 29 are summarized the known morphological char- acters of the known species of coccidia from rodents. Tables 1 through 21 give the characters of the species of Eimeria, arranged by host family or subfamily, while Tables 22 through 29 give the characters of the other genera of coccidia. 186 THE COCCIDIAN PARASITES OF RODENTS NUMBERS OF SPECIES OF RODENT CoOccIDIA A total of 196 named species of coccidia is included in this monograph. Eimeria is by far the most common genus, with 176 named species. In addition, 9 species of Isospora, 3 of Wenyonella, 2 each of Cryptosporid- zum and Klossiella, and 1 each of Dorisiella, Caryospora, Tyzzeria, and Klossia have been named. While this is quite a large number, it is only a small percentage of the number of species which must actually occur in rodents. In Table 30 are listed the numbers of genera and species in each rodent family together with the numbers of species of Eimeria which have been described from them. Eimeria has been described from only 15 per cent of the 337 genera and 4 per cent of the 2,688 species of rodents. Eimeria and other coccidia have been described from only 12 of the 24 rodent families listed. No coccidia have been named from such large rodent genera as Acomys (26 species), Dendromus (29 species), Otomys (26 spe- cies), Rhipidomys (27 species), Thomasomys (40 species), Akodon (62 species), Reithrodontomys (27 species), Gerbillus (52 species), Tatera (42 species), Pitymus (32 species), Cryptomys (46 species), Ctenomys (51 species), Coendou (25 species), Callosciurus (65 species), and Graphiurus (39 species). We know that many hosts have more than 1 species of Eimeria. For example, 8 have been described from Sciurus vulgaris, 5 from Apodemus sylvaticus, 8 from Mus musculus, 6 from Lophuromys sikapusi, and 6 from Myocastor coypus. On the other hand, we also know that some species of coccidia occur in more than | host. Assuming that there is an average of 1 Eimeria species per rodent species, we can estimate that there may be about 2,700 species of Eimeria alone in rodents. The num- ber already described is about 6.5 per cent of this. In addition, our information on most of the named species is meager. In most cases all that we have is a description of the oocysts, and many of these descriptions are incomplete. The location in the host is known for only 45 species of Eimeria (26 per cent of those named), the endog- enous stages are known for only 25 species (14 per cent of those named), and complete life cycles have been worked out for only 4 species of Eimeria (E. mohavensis, E. miyairu, E. nieschulz, and E. separata). RELATION OF EIMERIA OocysT MORPHOLOGY TO Host FAMILY In an attempt to determine whether there is any morphological char- acter or group of characters which might be used to differentiate the oocysts of Eimeria of one group of rodents from those of another group (or to differentiate the oocysts of rodent Eimerias from those of other DISCUSSION 187 host orders or classes), the known morphological characters of the oocysts of Eimeria species were tabulated by host family or subfamily in Table 31. In general, it can be said that the oocysts of rodent Eimeria species usually lack a micropyle (82 per cent of 176 species), lack an oocyst residuum (65 per cent of 171 species), may or may not have a polar granule (57 per cent of 156 species have one) or a sporocyst Stieda body (60 per cent of 126 species have one), and usually have a sporocyst re- siduum (89 per cent of 166 species). Rodent coccidia cannot be differentiated from the coccidia of other hosts on the basis of the presence or absence of any of these characters. A micropyle is more common in the Eimeria species from the Sciuridae (36 per cent of 42 species) than in those from other rodent families (13 per cent of 134 species). An oocyst polar granule is more common in the coccidia from the Muridae (66 per cent of 96 species) than in those from other rodent families (43 per cent of 60 species). Aside from these, there are no noteworthy differences in distribution of characters between Eimeria species from different host families. Certainly there is no evidence of parallel evolution between hosts and parasites among the rodent coccidia. The morphological characteristics of the coccidian species have not changed progressively, but appear to have arisen more or less at random. Cross-INFECTION STUDIES The cross-infection experiments which have been carried out with the species of Eimeria from rodents are summarized in Table 32. Such ex- periments have been carried out with only 20 species of Eimeria (12 per cent of the named species) from 16 species of donor hosts (17 per cent of the number from which Eimeria has been described). A total of 61 donor-receptor combinations was investigated, not including those in- volving subspecies of the same species. Of these, 56 were rodent-to-rodent, 3 rodent-to-lagomorph, and 2 rodent-to-carnivore. Of the rodent-to-rodent combinations, 9 were from one species to another in the same genus, while 47 were from one genus to another. Eight of the nine attempts to transfer Eimeria between species of the same genus were successful, but none of the attempts to transfer it to a species in a different genus, even within the same family, succeeded. While many more cross-infection studies should be carried out, those which have already been done indicate that the species of Ezmeria in rodents are highly host-specific. The only valid cross-infection experiment carried out with the other genera of rodent coccidia was an unsuccessful attempt to infect rats with Cryptosporidium muris from the mouse. 188 THE COCCIDIAN PARASITES OF RODENTS SUMMARY This monograph summarizes the known information on taxonomy, morphology, life cycle, hosts, location in the host, pathogenicity, geo- graphic distribution, and cross-transmission studies of the 196 named species of coccidia of rodents. These include 176 species of Eimeria, 9 ot Isospora, 3 of Wenyonella, 2 each of Cryptosporidium and Klossiella, and | each of Dorisiella, Caryospora, Tyzzeria, and Klossta. In addition, similar data are given for those forms for which insufficient information is available to justify assigning them names. Eimeria, which is the commonest genus, has been described from only 15 per cent of the 337 genera and 4 per cent of the 2,688 species of rodents. The location in the host is known for 45 species of Eimeria (26 per cent of those named), the endogenous stages are known for 25 species (14 per cent of those named), and complete life cycles have been worked out for only 4 species. Among the other coccidian genera, the location in the host, endogenous stages and life cycles are known only for Crypto- sporidium, Klossiella, and 1 species of Wenyonella. In general, the oocysts of rodent Eimeria usually lack a micropyle (82 per cent of 176 species in which the presence or absence of this character has been mentioned), lack an oocyst residuum (65 per cent of 171 species), may or may not have a polar granule (57 per cent of 156 species have one) or a sporocyst Stieda body (60 per cent of 126 species have one), and usually have a sporocyst residuum (89 per cent of 166 species). Rodent Eimerias cannot be differentiated from the Eimerias of other hosts on the basis of the presence or absence of any of these characters. No evidence of parallel evolution between hosts and parasites was ob- served among rodent Eimerias. The morphological characters of the parasite species have not changed progressively, but appear to have arisen more or less at random. Cross-infection experiments have been carried out with 20 species of Eimeria from 16 donor hosts. Eight out of 9 attempts to transfer Eimeria between species of the same rodent genus were successful, but none of 47 attempts to transfer it to a species in a different rodent genus, even within the same family, succeeded. The following new species of coccidia are described for the first time: Eimeria hansonorum n. sp. and FE. ferrist n. sp. from the wild house mouse (Mus musculus); and Isospora ratti n. sp. from the wild Norway rat (Rattus norvegicus). The following are established as new species of Eimeria: E. moelleri n. sp. for the form described from Sciurus carolinensis by MOller (1923) SUMMARY 189 and assigned by him to E. sciurorum; E. ascotensis n. sp. for the form described from Sciurus carolinensis by Webster (1960) and assigned by him to FE. neosciuri; E. kniplingi n. sp. for the form described from Sciurus niger rufwenter by Knipling and Becker (1935) and assigned by them to E. scturorum; E. asiatici n. sp. for the form described from Eutamias asiaticus by Tanabe and Okinami (1940) and assigned by them to E. beecheyi; E. batabatensis n. sp. for the form described from Arvicola terrestris by Musaev and Veisov (1960) and assigned by them to E. arvicolae; E. rysavyi n. sp. for the form described by RySavy (1957) from Clethrionomys glareolus and assigned by him to E. apodemi; FE. kazakhstanensis n. sp. for the form described from Ellobius talpinus by Svanbaev (1956) and assigned by him to E£. volgensis; E. talpini n. sp. for the form described from Ellobius talpinus by Svanbaev (1956) and assigned by him to E. beckert; E. tamariscini n. sp. for the form de- scribed from Meriones tamariscinus by Svanbaev (1956) and assigned by him to EF. musculi; E. hungaryensis n. sp. for the form described from Apodemus flavicollis by Pellérdy (1954) and assigned by him to E. muris; E. prasadi n. sp. for the form described from Apodemus sylvaticus by Svanbaev (1956) and assigned by him to EF. hindlez; E. svanbaevi n. sp. for the form described from Apodemus sylvaticus by Svanbaev (1956) and assigned by him to E. krijgsmanni; E. russiensis n. sp. for the form described from Apodemus sylvaticus by Svanbaev (1956) and assigned by him to E. musculi; E. heterocephali n. sp. for the form described from Heterocephalus glaber by Porter (1957) and as- signed by her to FE. muris; E. cernae n. sp. for the form described from Clethrionomys glareolus by Cerna (1962) and assigned by her to F. schueffneri; E. susliki n. sp. for the form described from Spermophilus maximus by Svanbaev (1962) and assigned by him to E. ussuriensis; E. peschankae n. sp. for the form described from Meriones tamariscinus by Svanbaev (1962) and assigned by him to E. krijgsmanni; E. assaensis n. sp. for the form described from Meriones tamariscinus by Svanbaev (1962) and assigned by him to E. callospermophili; and E. dorneyi Nn. sp. for the form described from Glaucomys sabrinus macrotis by Dorney (1962) and called Eimeria sp. by him. The following species of Eimeria are redescribed: E. nieschulzi from Rattus norvegicus in Mlinois and from R. hawaiiensis in Hawaii (new host record); E. separata from Rattus norvegicus in Illinois and from R., hawaiiensis in Hawaii (new host record); and E. falciformis from Mus musculus in I]linois. 190 LITERATURE CITED Allen, Ena A. 1936. Tyzzeria perniciosa gen. et sp. novy., a coccidium from the small intestine of the Pekin duck, Anas domesticus L. Arch. f. Protistenk. 87:262-267. Alves de Souza, M. 1931. Coccidiose em cobaya. Rev. Zootechn. Vet. 17:11-14. Andrews, J. M. 1927. Host-parasite specificity in the coccidia of mammals. J. Parasitol. 13:183-194. 1928. New species of coccidia from the skunk and prairie dog. J. Para- sitol. 14:193-194. Becker, E. R. 1934. Coccidia and coccidiosis of domesticated, game, and labora- tory animals and of man. Collegiate Press, Ames, Iowa, 147 pp. 1939. Effect of thiamin chloride on Eimeria nieschulz infection of the rat. Proc. Soc. Exper. 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Protozool. 8:134-138. Zolotarev, N. A. 1935. K voprosy o koktsidiyakh pyshnykh zverei. Sb. Rabot Dagestanskogo Protozoologicheskogo Nauch.-Issled. Opornogo Punkta Severo- kavkazskoi VOS i Narkomzema DASSR. 1935, vyp. 1. (Cited by Musaev and Veisov, 1959a). 1938. Zur Frage der Kokzidien in Zieseln. Wien. tierarztl. Monatschr. 25:658-661. Zwart, P., and W. J. Strik. 1961. Further observations on Eimeria dolichotis, a coccidium of the Patagonian cavy (Dolichotis patagonica). J. Protozool. 8:58-59. 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SUALOVAVH) ISAIOO SLNAGOU AO FPIYAZZAL 9g ATAVL (LI6I ‘Surpryog pue peqiiosap ION = snynanl snynovf uuewyj1ey) “ds (sistxa qsA900 and) OU EMEYORD A[qeqoid) TL) + repnsair] OF-0§ ‘SNpJa940d vIADD avikvqoo [eortayds 0} (sjstxa ysA000 ana) ou ATqeqoid) [eortaqdsqns Jajyaweip af = §1X<9I + Iepnsaqty 7 op 01 dq snpNIsNU sn srinul wan aaog (suOIOTJN) Wan aATANVYD (suOIDTIN) SaIOddS -GIsay VaalLg AZIS -aIsdy AVIOG ATAMONIST TIVM AdVHS AZIS SISOH D1) a1SSO] ¥ SUTLOVAVH) LSXDOUOdS SUALOVAVH') ISADOO SLNAGOUW AO PTTHISSOTY 6g ATAV.L yoryy 7 GO ‘umorq ayed 19uUT [eorsayds sprya 7 Z ‘yOoUTS ESL ‘Juatedsueszy ‘ssa[10 OF X8F IN snyojnaiupUt AP = FI-SI = — -[09 193nNO :sI9hP] Z [eptosdi[q FF-GE X §o-SF snasKwmoslad suaxalqiad UE EE Soe SS 07 OA et ee LL Ne a ae el ee ee wan adog (suOoIDTJ) wan aTANVAD (suOAITJA) SaIOAdS -GISHY + VWAaILS AZIS -isay AVIOG AIAdOAIIY TIVM AdVHS AZIS sISOH DISSO]M SUALOVUVH') ISKOOUOdS SUALOVUVH’) ISADOQ om a re en ee 8868686068086 8685680858008 8 SLNAGOU AO VISSOTN gg Wav TABLE 30 NAMED SPECIES OF EIMERIA IN RODENTS No. Eimeria No. No. SPECIES RODENT RODENT DE- RODENT FAMILY GENERA* SPECIES* SCRIBED Bathyergidae 5 53 1 Echimyidae 28 152 6 Dinomyidae 1 1 0 Erethizontidae 4 29 0 Dasyproctidae 2 29 4 Hystricidae 4 28 0 Cuniculidae ] 2 0 Chinchillidae 3 d 0 Caviidae 5 22 2, Hydrochoeridae ] 2 2 Aplodontiidae 1 1 0 Sciuridae 44 386 39 Castoridae 1 4 ] Heteromyidae 5 103 6 Geomyidae 9 96 2 Anomaluridae 4 9 0 Pedetidae 1 2 0 Ctenodactylidae 4 8 0 Dipodidae 15 50 3 Muscardinidae 9 54 4 Lophiomyidae ] 1 0 Spalacidae l 8 0 Rhizomyidae 2 8 0 Muridae 186 1633 106 (Subf. Murinae) (71) (608) (37) (Subf. Rhynchomyinae) (1) (1) 0 (Subf. Hydromyinae) (7) (8) 0 (Subf. Dendromyinae) — (5) (52) 0 (Subf. Deomyinae) (1) (1) 0 (Subf. Otomyinae) (2) (28) 0 (Subf. Cricetinae) (54) (499) (25) (Subf. Gymnuromyinae) (1) (1) 0 (Subf. Tachyoryctinae) (2) (16) (1) (Subf. Gerbillinae) (12) (145) (28) (Subf. Myospalacinae) (1) (13) 0 (Subf. Microtinae) (29) (261) (15) Total 337 2688 176 RODENT GENERA FROM WHICH Eimeria Has BEEN DESCRIBED RODENT SPECIES FROM WHICH Eimeria Has BEEN DESCRIBED Per No. Cent 1 20 1 4 0 0 2 100 0 0 0 2 40 ] 100 0 10 23 ] 100 3 60 2 22 0 0 0 ] 7 3 33 0 0 0 25 13 @ (13) 0 0 0 0 0 (8) (15) 0 (1) (50) (1) (8) 0 (6) Ql) 52 15 Per No. Cent 1 2 1 1 0 0 2, a 0 0 0 3 14 1 50 0 26 7 1 25 U 7 2 2 0 0 0 2 4 3 6 0 0 0 46 3 (15) 2) 0 0 0 0 0 (15) (8) 0 (ly 6) (6) (4) 0 9) (3) 95 a * Based on a count of genera and species listed by Ellerman, J. R. 1940-1941. genera of living rodents. London, Brit. Mus. (Nat. Hist.). 9 4 The families and vol., xxvi + 689, xii + 690 pp. IT 68 OF 09 ¢9 CE €F LG 68 81 49] IDADYI YyIva LOf ]D}0} fo Juad sag 81 SFI 1g cL nal 09 L9 68 FFI of 1P10.]. 0) I I 0 oepisiadyieg 0 9 § @ 9 0 g @ P G oeprAuryoy 0 § 0 G § 0 § 0 § 0 aepro0iddseq 0 G 0 } @ 0 és 0 6 0 9ePlI20yIoIpAH 0 3 0 I I I 3 0 3 0 JepIARy 6 I 6 I é I § 0 ovpipodiq, 0 ¢ 3 I g 0 I @ é G oeprurpieosn jy (0) (1) (0) (1) (1) (0) aeunoXrod yor TF, (¢ (16 (¢) (€1) (0g) (F) (On) ree) (38) (g) aeULIN |, (F) (£3 (£1) (9) (FI) (FT) (Or) (L1) (F3) (g) aeury[Iq.i9a5) (6 (1) (¢) (¢) (6) (¢) (9) (9) (11) (F) IeUTIOINT IN (1) (F3) (¢) (81) (11) (L1) (L) (81) (F3) (3) IeUTIIIIAD él 68 rae oF £9 IF 6 69 66 ral aeplinyy 0 0 I 0 0 I 0 I I 0 IEPIIO}se-) 0 9 I G 3 F g g 9 0 aeprdwo.19j9H 0 3 I I ra 0 i 0 I I aeprduloasy 7 0g II 61 93 All 91 LI LE cI depLINog a ee ee eee a = = tae = ail = = = IP ATINV LNAGOY WANGISAY LSADOUOdS AGOG VQuILsg WONGISAY LISXDOQO ATONVUS‘) UVIOd WIACOWI]N SUTLOVAVH) GALVNOISA(T LNOHLIM YO HLM Saloddg visaway sO Suda N Se Ee ee ee Se Se ee SAVTINVA LNAGOW LNAWATMIG WOU SAIOAdS PINAWIT LO SLSADOO AO SOLLSTARLOVUVHD TWOIOMOTOHAUOWN I§ ATAV.T, TABLE 32 SUMMARY OF CROSS-INFECTION EXPERIMENTS WITH EIMERIA IN RODENTS Eimeria SPECIES Donor Host RECEPTOR HOST RESULT REFERENCE sciurorum mira moelleri cynomysis citelli petauristae mohavensis leucopi tachyoryctis apionodes Sciurus vulgaris Sciurus vulgaris Sciurus carolinensis Cynomys ludovicianus Spermophilus tri- decemlineatus Petaurista inornatus Dipodomys panamin- tinus mohavensis Peromyscus leucopus Tachyoryctes ruandae A podemus flavicollis Rattus norvegicus Spermophilus citellus Glis glis European domestic squirrel (Sciurus vulgaris ?) Domestic cat Rattus norvegicus Mus musculus Indian rabbit Dipodomys panamintinus leucogenys Dipodomys p. panamintinus Dipodomys p. caudatus Dipodomys m. merriami Dipodomys brevinasus Dipodomys heermanni morroensts Dipodomys h, tularensis Dipodomys h. swarthi Dipodomys d. deserti Dipodomys a. agilis Perognathus longimembris Perognathus formosus mohavensis Peromyscus boylii Peromyscus maniculatus Peromyscus californicus Peromyscus truei Onychomys torridus Neotoma lepida Spermophilus leucurus Mus musculus Rattus norvegicus Peromyscus nuttalli Rattus norvegicus Mus musculus Mus musculus Microtus arvalis Clethrionomys glareolus Cricetus cricetus (ete }++++ 4+4+4+4++ 4+ | Galli-Valerio, 1922 Pellérdy, 1954 Moller, 1923 Andrews, 1927 Kartchner and Becker, 1930 Ray and Singh, 1950 Doran, 1951, 1953 Von Zellen, 1961 van den Berghe and Chardome, 1956 Pellérdy, 1954 TABLE 32 (Continued) Eimeria SPECIES Donor Host RECEPTOR Host RESULT REFERENCE apodemi rugosa hungaryensis vinckei nieschulzi falciformis oryzomysi phyllotis weissi dolichotis Apodemus flavicollis Apodemus flavicollis Apodemus flavicollis Thamnomys s. sur- daster Rattus rattus Rattus norvegicus Rattus norvegicus Mus musculus Mus musculus Oryzomys sp. Phyllotis a. amicus (?) Phyllotis a. amicus (?) Mus musculus Microtus arvalis Clethrionomys glareolus Cricetus cricetus Mus musculus Microtus arvalis Clethrionomys glareolus Cricetus cricetus Mus musculus Microtus arvalis Clethrionomys glareolus Cricetus cricetus Rattus norvegicus Mus musculus Rattus norvegicus Rattus rattus Mus musculus Cavia porcellus Oryctolagus cuniculus Mus musculus Oryctolagus cuniculus Rattus norvegicus Canis familiaris Rattus norvegicus Mus musculus Mus musculus Rattus norvegicus Mus musculus Cavia porcellus Dolichotis patagonum Cavia porcellus Pellérdy, 1954 Pellérdy, 1954 Pellérdy, 1954 Rodhain, 1954 Dieben, 1924 Dieben, 1924 Pérard, 1926 Noller, 1920 Pérard, 1926 Carini, 1935 Gonzales-Muga- buru, 1942 Gonzales-Muga- buru, 1946 Zwart and Strik, 1961 « om) ‘ « at ‘ : P y — i, € 3 ot iia 7 i ~ a “ 2 ~~ 7 T a - ae _* j : s t a $ ! 7 = eee . : baa _ _ on y - = - . 2 4 +4 - = ‘ | — am . Oo a” a » _ > ( i |—CURe 7 4 ooh : a 7 4 i a a 7 7 - +, Xe _ ‘ eee ay) - » PLATES 254 PLATE | Fics. 1-4. Eimeria sciurorum Galli-Valerio, 1922. Oocysts from Scturus vulgaris (from Yakimoff, Sokoloff, and Rastegaieff, 1931). < 1300. Fics. 5 ANp 6. Eimeria serbica Pop-Cenitch and Bordjochki, 1957. Oocysts from Sciurus vulgaris (?) (from Pop-Cenitch and Bordjochki, 1957). < 1300. Fics. 7 ANpD 8. Eimeria neosciuri Prasad, 1960. Oocysts from Sciurus carolinensis (from Prasad, 1960). x 1700. Fic. 9. Eimeria neosciuri Prasad, 1960. Oocyst from Scturus vulgaris (from RySavy, 1954). « 1700. Fics. 10-14. Eimeria neosciuri Prasad, 1960 from Sciwrus carolinensis (from Prasad, 1960). « 900. 10. Young schizont in intestinal cell. 11. Schizont containing 13 nuclei in intestinal cell. 12. Schizont containing merozoites in intestinal cell. 13, 14. Developing macrogametes in intestinal cells. Fics. PLATE 2 15 ano 16. Eimeria neosciuri Prasad, 1960 from Sciurus carolinensis (from Prasad, 1960). « 900. 15. 16. Mature macrogamete showing plastic granules. Mature microgametocyte containing microgametes. Fics. 17-24. Eimeria ascotensis n. sp. from Sciurus carolinensis (from Web- ster, 1960). « 2600. live 18. 19. 20. “alle 22, 2a. Dee Undifferentiated young stages in intestinal cells. Schizont containing merozoites. Endogenous oocyst. Microgametocyte containing microgametes. Mature macrogamete. Unsporulated oocyst. Developing oocyst. Sporulated oocyst. no Or ie.) PLATE 3 Fics. 25-27. Eimeria kniplingi n. sp. Oocysts from Sciurus niger rufiwenter (from Knipling and Becker, 1935). 2200. Fic. 28. Eimeria sp. Bond and Bovee, 1957. Oocyst from Sciurus carolinen- sis (from Bond and Bovee, 1957). « 1400. Fic. 29. Eimeria tamiasciurt Levine, Ivens, and Kruidenier, 1957. Oocyst from Tamiasciurus hudsonicus (from Levine, Ivens, and Kruidenier, 1957). x 2650. Fic. 30. Eimeria toddi Dorney, 1962. Oocyst from Tamiasciurus hudsonicus (from Dorney, 1962). 1200. Fics. 31-35. Eimeria garnhami McMillan, 1958 from Xerus erythropus (from McMillan, 1958). 31. Mature schizont with merozoites. « 600. 32. Nearly mature microgametocyte. « 1800. 33. Nearly mature macrogamete. & 1700. 34. Intracellular zygote. No measurements. 35. Undifferentiated oocyst from fresh feces. &« 1900. (=>) Nt PLATE 4 Fics. 36 ann 37. Eimeria garnhami McMillan, 1958 from Xerus erythropus (from McMillan, 1958). x 1800. 36. Undifferentiated oocyst 90 days old. 37. Sporulated oocyst. Fics. 38 AND 39. Eimeria monacis Fish, 1930. Oocysts from Marmota monax (from Fish, 1930). < 2000. Fic. 40. Eimeria cynomysis Andrews, 1928. Oocyst from Cynomys ludovt- cianus (from Andrews, 1928). 1500. Fics. 41-44. Eimeria citelli Kartchner and Becker, 1930 from Spermophilus tridecemlineatus (from Kartchner and Becker, 1930). « 2040. 41. Unsegmented oocyst from fresh feces. 42. Oocyst with sporocyst prior to formation of sporozoites but showing refractile bodies. 43. Sporulated oocyst. 44. Section of an infected cecal wall showing gametes and gametocytes within cells of glands. Fic. 45. Eimeria glaucomydis Roudabush, 1937. Oocyst from Glaucomys volans (from Roudabush, 1937). « 1900. 261 wee \\ Wis \ Dyas) SLO Oke nC BNC Ce Dt: ws a: ay 7% Ore SlinllinuiiZ ea iT I \ak PLATE 5 Fics. 46-48. Eimeria citelli Kartchner and Becker, 1930 from Spermophilus pygmaeus (from Sassuchin and Rauschenbach, 1932). « 1800. 46. Unsporulated oocyst. 47. Oocyst showing incomplete sporulation. 48. Sporulated oocyst. Fic. 49. Eimeria bilamellata Henry, 1932. Oocyst from Spermophilus franklinii (from Hall and Knipling, 1935). « 2000. Fic. 50. Eimeria volgensis Sassuchin and Rauschenbach, 1932. Oocyst from Spermophilus pygmaeus (from Sassuchin and Rauschenbach, 1932). « 1800. Fic. 51. Eimeria franklinii Hall and Knipling, 1935. Oocyst from Spermophilus franklinn (from Hall and Knipling, 1935). & 2000. Fics. 52. anp 53. Eimeria hoffmeisteri Levine, Ivens, and Kruidenier, 1958. Oocysts from Spermophilus spilosoma. 52. From Levine, Ivens, and Kruidenier, 1958. * 2600. 53. From Ivens, Kruidenier, and Levine, 1959. « 2600. 263 264 PLATE 6 Fic. 54. Eimeria callospermophili Henry, 1932. Oocyst from Spermophilus spilosoma (from Levine, Ivens, and Kruidenier, 1958). « 2600. Fic. 55. Eimeria vilast Dorney, 1962. Oocyst from Tamias striatus (from Dorney, 1962). « 2600. Fic. 56. Eimeria wisconsinensis Dorney, 1962. Oocyst from Tamuas striatus (from Dorney, 1962). x 1200. Fic. 57. Eimeria parasciurorum Bond and Bovee, 1957. Oocyst from Glau- comys volans (from Roudabush, 1937). « 1600. Fic. 58. Eimeria lateralis Levine, Ivens, and Kruidenier, 1957. Oocyst from Spermophilus lateralis (from Levine, Ivens, and Kruidenier, 1957). X 2650. 265 PLATE 7 Fic. 59. Eimeria eutamiae Levine, Ivens, and Kruidenier, 1957. Oocyst from Eutamias dorsalis (from Levine, Ivens, and Kruidenier, 1957). > 2650. Fic. 60. Eimeria geomydis Skidmore, 1929. Oocyst from Geomys bursarius (from Skidmore, 1929). < 2100. Fics. 61-65. Eimeria parasciurorum Bond and Bovee, 1958 from Glaucomys volans (from Bond and Bovee, 1958). & 1300. 61. Unsporulated oocyst. 62. Unsporulated oocyst prior to segmentation. 63. Oocyst with sporoblasts and polar body which quickly disintegrates. 64. Oocyst showing immature sporocysts with developing sporozoites. 65. Completely sporulated oocyst. Fics. 66-69. Eimeria petauristae Ray and Singh, 1950 from Petaurista pe- taurista (from Ray and Singh, 1950). « 1100. 66. Sporulated oocyst with outer covering ruptured. 67. Sporocyst liberated from the oocyst. 68. Sporozoite coming out of narrow end of sporocyst. 69. Liberated sporozoite. 267 268 PLATE 8 Fic. 70. Eimeria thomomysis Levine, Ivens, and Kruidenier, 1957. Oocyst from Thomomys bottae (from Levine, Ivens, and Kruidenier, 1957). « 2650. Fic. 71. Eimeria perognathi Levine, Ivens, and Kruidenier, 1957. Oocyst from Perognathus intermedius (from Levine, Ivens, and Kruidenier, 1957). 2650. Fics. 72 anp 73. Eimeria mohavensis Doran and Jahn, 1949 from Dipodomys mohavensis (from Doran and Jahn, 1952). « 1700. 72. Immature oocyst showing two polar bodies. 73. Completely sporulated oocyst. Fics. 74-77. Eimeria mohavensis Doran and Jahn, 1949 from Dipodomys p. mohavensis and D. merriami (from Doran, 1953). 74. Mature macrogamete. & 2500. 75. Microgametocyte. < 2300. 76. Merozoite packet (small type). « 3200. 77. Merozoite packet (large type). & 3200. Fic. 78. Eimeria penicillati Ivens, Kruidenier, and Levine, 1959. Odocyst from Perognathus penicillatus (from Ivens, Kruidenier, and Levine, 1959). < 2600. 269 Pr a pea O os * p . On SO ee Oor eG we Siete OR a a 270 PLATE 9 Fics. 79 AND 80. Eimeria mohavensis Doran and Jahn, 1949 from Dipodomys p. mohavensis (from Doran and Jahn, 1952). 79. Section through cecum. MA-Macrogamete; MI-Microgametocyte; SCH- Schizont. 80. Diagram of intestinal tract showing locations of schizogony. SI-Small Intestine; CA-Cecum. no — ho ~I ho PLATE 10 Fic. 81. Eimeria dipodomysis Levine, Ivens, and Kruidenier, 1958. Oocyst from Dipodomys phillips: (from Levine, Ivens, and Kruidenier, 1958). x 1400. Fic. 82. Eimeria couesit Kruidenier, Levine, and Ivens, 1960. Oocyst from Oryzomys couest (from Kruidenier, Levine, and Ivens, 1960). x 2650. Fic. 83. Eimeria liomysis Levine, Ivens, and Kruidenier, 1958. Oocyst from Liomys pictus (from Levine, Ivens, and Kruidenier, 1959). x 2600. Fic. 84. Eimeria picti Levine, Ivens, and Kruidenier, 1958. Oocyst from Liomys pictus (from Levine, Ivens, and Kruidenier, 1958). x 2600. 273 274 PLATE 11 Fic. 85. Eimeria peromysci Levine, Ivens, and Kruidenier, 1957. Oocyst from Peromyscus truei (from Levine, Ivens, and Kruidenier, 1957). x 2650. Fic. 86. Eimeria langebarteli Ivens, Kruidenier, and Levine, 1959. Oocyst from Peromyscus boylii (from Ivens, Kruidenier, and Levine, 1959). x 2650. Fics. 87 AND 88. Eimeria arizonensis Levine, Ivens, and Kruidenier, 1957 (from Levine and Ivens, 1960.) & 2600. 87. Oocyst from Peromyscus leucopus. 88. Oocyst from Peromyscus maniculatus. & wy lon | Af ‘ e ~ no ~I ee) PLATE 13 Fics. 94-97. Eimeria leucopi von Zellen, 1961 from Peromyscus leucopus (from von Zellen, 1961). 2650. 94. Newly discharged cyst after protoplasmic withdrawal. 95. Clear area subsequent to nuclear disappearance. 96. Buckelbildung or ball stage. Refractile globules associated with mound formation. 97. Buckelbildung or ball stage. Formation of four mounds. 280 PLATE 14 Fics. 98-101. Eimeria leucopi von Zellen, 1961 from Peromyscus leucopus (from von Zellen, 1961). « 2650. 98. Sporoblasts, with clear polar caps, separate from granular residuum. 99. Pyramid stage. Iwo days, fourteen hours. 100. Late pyramid stage. ‘Two hours after Fig. 99. 101. Sporoblast elongation. Globular material retained within sporoblasts. 281 PLATE 15 Fic. 102. Eimeria leucopi von Zellen, 1961. Completely sporulated oocyst from Peromyscus leucopus (from von Zellen, 1961). 3800. Fic. 103. Eimeria baiomysis Levine, Ivens, and Kruidenier, 1958. Oocyst from Baiomys taylori (from Levine, Ivens, and Kruidenier, 1958). > 2600. Fic. 104. Ezmeria onychomysis Levine, Ivens, and Kruidenier, 1957. Oocyst from Onychomys leucogaster (from Levine, Ivens, and Kruidenier, 1957). « 2650. Fic. 105. Eimeria albigulae Levine, Ivens, and Kruidenier, 1957. Oocyst from Neotoma albigula (from Levine, Ivens, and Kruidenier, 1957). « 2650. 284 PLATE 16 Fic. 106. Eimeria davisi Ivens, Kruidenier, and Levine, 1959. Oocyst from Neotoma albigula (from Ivens, Kruidenier, and Levine, 1959). 2600. Fic. 107. Eimeria migratoria Musaey and Veisov, 1961. Oocyst from Cricetulus migratorius (from Musaev and Veisov, 1961). « 2000. Fic. 108. Eimeria cricetuli Musaev and Veisov, 1961. Oocyst from Cricetulus migratorius (from Musaev and Veisov, 1961). « 2000. Fics. 109 ANp 110. Eimeria jardimlinica Musaev and Veisov, 1961. Oocysts from Cricetulus migratorius (from Musaevy and Veisov, 1961). & 2000. 286 PLATE 17 Fics. 111 anp 112. Eimeria immodulata Musaev and Veisov, 1961. Oocysts from Cricetulus migratorius (from Musaev and Veisov, 1961). x 2000. Fic. 113. Eimeria arusica Musaev and Veisov, 1961. Oocyst from Cricetulus migratorius (from Musaev and Veisov, 1961). « 2000. Fic. 114. Eimeria microtina Musaev and Veisov, 1959. Oocyst from Microtus socialis (from Musaev and Veisov, 1959). > 2000. Fics. 115 ANp 116. Eimeria wenricht Saxe, Levine, and Ivens, 1960. Oocysts from Microtus pennsylvanicus (from Saxe, Levine, and Ivens, 1960). 2580. Fics. 117 anp 118. Eimeria bohemica RySavy, 1957 from Arvicola terrestris (from RySavy, 1957). « 1800. 117. Unsporulated oocyst. 118. Sporulated oocyst. Fic. 119. Eimeria batabatensis n. sp. Oocyst from Arvicola terrestris (from Musaev and Veisov, 1960). & 2000. Fic. 120. Eimeria dicrostonicis Levine, 1952. Oocyst from Dicrostonyx groenlandicus richardsoni (from Levine, 1952). « 2300. > by Oo/ ) PLATE 18 Fic. 121. Eimeria terrestris Musaev and Veisoy, 1960. Oocyst from Arvicola terrestris (from Musaev and Veisov, 1960). « 2000. Fic. 122. Eimeria talischaensis Musaev and Veisov, 1960. Oocyst from Arvicola terrestris (from Musaev and Veisov, 1960). > 1600. Fics. 123-125. Eimeria rysavyi n. sp. from Clethrionomys glareolus. 123. Oocyst (from RySavy, 1957). « 1600. 124, 125. Oocysts (from Cerna, 1962). « 1440. Fic. 126. Eimeria noraschenica Musaev and Veisov, 1960. Oocyst from Meriones persicus (from Musaev and Veisov, 1960). * 1700. Fics. 127 aANnp 128. Eimeria disaensis Musaev and Veisov, 1960. Oocysts from Meriones persicus (from Musaev and Veisov, 1960). & 1400. Fics. 129 ann 130. Eimeria lerikaensis Musaev and Veisov, 1960. Oocysts from Meriones persicus (from Musaev and Veisov, 1960). « 1800. Fic. 131. Eimeria salasuzica Musaeyv and Veisov, 1960. Oocyst from Meriones persicus (from Musaey and Veisov, 1960). « 2100. PLATE 19 Fic. 132. Eimeria cernae n. sp. Oocyst from Clethrionomys glareolus (from Cerna, 1962). « 2900. Fic. 133. Eimeria schachtachtiana Musaev and Veisoy, 1960. Oocyst from Meriones shawi (from Musaev and Veisov, 1960). & 2000. Fic. 134. Eimeria nehramaensis Musaey and Veisov, 1960. Oocyst from Meriones shawi (from Musaevy and Veisov, 1960). « 2100. Fic. 135. Eimeria dzhahriana Musaev and Veisov, 1960. Oocyst from Meriones shawi (from Musaev and Veisov, 1960). « 2000. Fic. 136. Eimeria araxena Musaev and Veisov, 1960. Oocyst from Meriones shawi (from Musaev and Veisov, 1960). x 2800. Fic. 137. Eimeria astrachanbazarica Musaev and Veisov, 1960. Oocyst from Meriones shawi (from Musaev and Veisov, 1960). « 2500. Fic. 138. Eimeria zulfiaensis Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). « 1700. 291 PLATE 20 Fic. 139. Eimeria vinogradovi Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). « 2900. Fic. 140. Eimeria jurschuaensis Veisov, 1961. Oocyst from Meriones vino- gradovi (from Veisov, 1961). & 2300. Fic. 141. Eimeria sadaraktica Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). « 1800. Fic. 142. Eimeria bistratum Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). « 2600. Fic. 143. Eimeria arabiana Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). 2700. 293 | TA Leia PLATE 2] Fic. 144. Eimeria tasakendica Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). « 2800. Fic. 145. Eimeria musajevi Veisov, 1961. Oocyst from Meriones vinogradovi (from Veisov, 1961). & 2300. Fic. 146. Eimeria erythrourica Musaev and Alieva, 1961. Oocyst from Meriones libycus (from Musaev and Alieva, 1961). > 2000. Fic. 147. Eimeria schamchorica Musaey and Alieva, 1961. Oocyst from Meriones libycus (from Musaev and Alieva, 1961). « 1600. Fic. 148. Eimeria achburunica Musaey and Alieva, 1961. Oocyst from Meriones libycus (from Musaev and Alieva, 1961). & 1500. Fic. 149. Eimeria sumgaitica Musaev and Alieva, 1961. Oocyst from Meriones libycus (from Musaev and Alieva, 1961). & 2000. Fic. 150. Eimeria martunica Musaev and Alieva, 1961. Oocyst from Meriones libycus (from Musaev and Alieva, 1961). 1700. 0 ivy) 296 PLATE 22 Fic. 151. Eimeria apionodes Pellérdy, 1954. Oocyst from Apodemus flavicollis (from Pellérdy, 1954). 2100. Fics. 152 anp 153. Eimeria apodemi Pellérdy, 1954. 152. Oocyst from Apodemus flavicollis (from Pellérdy, 1954). * 2100. 153. Oocyst from A. flavicollis and A. sylvaticus (from Cerna, 1962). x 1500. Fic. 154. Eimeria poljanskii Veisov, 1961 emend. Oocyst from Meriones vinogradovi (from Veisov, 1961). 1900. Fics. 155 anp 156. Eimeria hungaryensis n. sp. 155. Oocyst from A podemus flavicollis (from Pellérdy, 1954). > 2000. 156. Oocyst from Apodemus sylvaticus (from Cerna, 1962). < 2200. Fic. 157. Eimeria tachyoryctis van den Berghe and Chardome, 1956. Oocyst from Tachyoryctes ruandae (from van den Berghe and Chardome, 1956). 3000. . = 5 orseveve~ Ss 6 eo, p 20% OG RA PLATE 23 Fic. 158. Eimeria rugosa Pellérdy, 1954. Oocyst from Apodemus flavicollis (from Pellérdy, 1954). & 2100. Fic. 159. Eimeria svanbaevi n. sp. Oocyst from Apodemus sylvaticus (from RySavy, 1954). x 1400. Fics. 160 anp 161. Eimeria sp. (RySavy, 1954) syn. Eimeria keilint Yakimoft and Gousseff of RySavy, 1954 and of Cerna, 1962. 160. Oocyst from Apodemus sylvaticus (from RySavy, 1954). & 1500. 161. Oocyst from Apodemus flavicollis (from Cerna, 1962). < 1500. Fic. 162. Eimeria dasymysis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Dasymys incomptus rufulus (from Levine, Bray, Ivens, and Gunders, 1959). x 2600. Fic. 163. Eimeria lemniscomysis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lemniscomys striatus striatus (from Levine, Bray, Ivens, and Gunders, 1959). > 2600. Fic. 164. Eimeria putevelata Bray, 1958. Oocyst from Lemniscomys striatus striatus (from Bray, 1958). < 1600. Fics. 165 anp 166. Eimeria miyairti Ohira, 1912. 165. Oocyst from Rattus norvegicus (from Becker, 1934). x 1200. 166. Oocyst from Rattus norvegicus and R. rattus (?) (from Matubayasi, 1938). « 1200. 299 300 PLATE 24 Fics. 167-181. Eimeria miyairii Ohira, 1912. 167. Oocysts from Rattus norvegicus (from Pinto, 1928). x 4000. 168-181. From Rattus norvegicus (from Roudabush, 1937). 168. Sporozoite. « 2300. 169. Sporozoite in host cell. & 2300. 170. Group of first-generation merozoites. & 1500. 171. First-generation merozoite. < 1800. 172. Schizont (probably second-generation). 173. Group of second-generation merozoites. 1900. 174. Second-generation merozoite. & 2500. 175. Group of third-generation merozoites. < 1400. 176. ‘Third-generation merozoite. & 2300. 177. Microgametocyte (surface view). 178. Microgamete. « 3000. 179. Double infection with young macrogametes shown below epithelium. 180. Macrogamete. 181. Figure copied by Roudabush from Ohira (1912) showing typical mac- rogamete. 301 PLATE 25 Fics. 182-187. Eimeria nieschulzi Dieben, 1924. 182. Oocyst from Rattus norvegicus (from Pérard, 1926). « 2000. 183, 184. Oocysts from R. norvegicus and R. rattus (?) (from Matubayasi, 1938). « 1400. 185. Oocyst from R. norvegicus (from Becker, 1934). & 1100. 186. Oocyst from R. norvegicus (from Becker, Hall, and Hager, 1932). x 2200. 187. Oocyst from R. norvegicus (original). 2600. Fics. 188-193. Eimeria separata Becker and Hall, 1931. 188. Oocyst from Rattus norvegicus (from Becker, Hall, and Hager, 1932). x 1800. 189. Oocyst from R. norvegicus (from RySavy, 1954). « 2400. 190. Oocyst from R. norvegicus (from Becker, 1934). « 1100. 191. Oocyst from R. norvegicus (original). « 2600. 192. Oocyst from R. hawaziensis (original). x 2600. 193. Oocyst from Rattus (Dephomys) defua (from Levine, Bray, Ivens, and Gunders, 1959). « 2600. 303 304 PLATE 26 Fics. 194-208. Eimeria nieschulzi Dieben, 1924 from Rattus norvegicus (from Roudabush, 1937). 194. LOD: 196. Sporozoite. « 2400. Sporozoite with divided posterior globule. > 2400. Cross section of first-generation merozoites in cell; note refractile globule. NOs 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. Group of first-generation merozoites. < 1900. First-generation merozoite. « 2300. Group of second-generation merozoites. < 1900. Second-generation merozoite. & 2600. Group of third-generation merozoites. « 2100. ‘Third-generation merozoite. & 2100. Group of fourth-generation merozoites. & 1800. Fourth-generation merozoite. & 2500. Developing microgametocyte. Microgametocyte (surface view). Mature macrogamete. Microgamete. « 1800. 305 306 PLATE 27 Fics. 209-214. Eimeria nieschulzi Dieben, 1924 from Rattus norvegicus (from Roudabush, 1937). Development of the oocyst wall. Plastic granules shown as white bodies with a solid outline. Haematoxylinophilic granules shown as large black dots. 209. Early development showing migration of granules. 210. Plastic granules have almost reached the periphery. Second group remains close to the nucleus. 211. Plastic granules have reached the edge and have begun to flatten out. Haematoxylinophilic granules have begun to elongate. 212. Outer wall formed from plastic granules. Inner wall forming from haematoxylinophilic granules. 213. Inner wall almost fully formed. 214. Mature oocyst inner wall shrunken away from outer wall. Second set of plastic granules shown in cytoplasm. Fics. 215-227. Eimeria separata Becker and Hall, 1931 from Rattus norvegicus (from Roudabush, 1937). 215. Sporozoite. & 2400. 216. Sporozoite after rounding up. 217. Early first-generation merozoites. 218. Group of first-generation merozoites. « 1600. 219. First-generation merozoite. & 2300. 220. Early second-generation schizont (double infection). 221. Group of second-generation merozoites. < 1700. 222. Second-generation merozoite. & 2500. 223. Group of third-generation merozoites. < 1900. 224. ‘Third-generation merozoite. & 2300. 225. Microgametocyte (surface view). 226. Almost mature macrogamete. 227. Microgamete. & 2400. 307 308 PLATE 28 Fic. 228. Eimeria praomysis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Rattus (Praomys) tullbergi rostratus (from Levine, Bray, Ivens, and Gunders, 1959). « 2600. Fics. 229-240. Eimeria falciformis (Eimer, 1870) Schneider, 1875 from Mus musculus. 229. Sporulated oocyst (original). « 2700. 230, 231. Sporulated oocyst and sporozoites, respectively (from Reichenow, 1953). 232-239. From Wenyon (1926). « 2400. 232, 233, 234, 235. Bundles of merozoites. 236 and 237. Sections through bundles of merozoites. 238. Microgametocyte. 239. Macrogamete. 240. Sporulated oocyst (from Cerna, 1962). & 2500. Fic. 241. Eimeria hansonorum n. sp. Oocyst from Mus musculus (original). « 2600. 309 310 PLATE 29 Fic. 242. Eimeria ferrisi n. sp. Oocyst from Mus musculus (original). X 2600. Fic. 243. Eimeria musculoidei Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Mus (Leggada) musculoides (from Levine, Bray, Ivens, and Gunders, 1959). x 2600. Fic. 244. Eimeria lophuromysis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. sikapusi (from Levine, Bray, Ivens, and Gunders, 1959). x 2600. Fic. 245. Eimeria sikapusii Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. sikapusi (from Levine, Bray, Ivens, and Gunders, 1959). x 2600. Fic. 246. Eimeria liberiensis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. sikapusi (from Levine, Bray, Ivens, and Gunders, 1959). x 2600. Fic. 247. Eimeria africana Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. sikapusi (from Levine, Bray, Ivens, and Gunders, 1959). < 2600. 311 312 PLATE 30 Fic. 248. Eimeria harbelensis Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. sikapusi (from Levine, Bray, Ivens, and Gunders, 1959). S< 2600. Fic. 249. Eimeria schoutedeni van den Berghe and Chardome, 1957. Oocyst from Cricetomys dissimilis (from van den Berghe and Chardome, 1957). & 3500. Fic. 250. Eimeria dyromidis Zolotarev, 1935. Oocyst from Dryomys nitedula (from Musaev and Veisov, 1959). « 2700. Fic. 251. Eimeria nachitschevanica Musaev and Veisov, 1959. Oocyst from Dryomys nitedula (from Musaey and Veisov, 1959). « 2600. Fics. 252 and 253. Eimeria dolichotis Morini, Boero, and Rodriguez, 1955 from Dolichotis patagonum patagonum (from Morini, Boero, and Rodriguez, P15), 252. Sporulated oocyst. & 1700. 253. Sporocyst. < 3000. 313 314 PLATE 31 Fic. 254. Eimeria caviae Sheather, 1924. Oocyst from Cavia porcellus (from RySavy, 1954). x 1500. Fic. 255. Eimeria myopotami Yakimoff, 1933. Oocyst from Myocastor coypus (from Prasad, 1960). > 1300. Fics. 256 aAnp 257. Eimeria nutriae Prasad, 1960. Oocysts from Myocastor coy pus (from Prasad, 1960). « 1300. Fic. 258. Eimeria myocastori Prasad, 1960. Oocyst from Myocastor coypus (from Prasad, 1960). « 1600. Fic. 259. Eimeria kruidenieri Levine, Bray, Ivens, and Gunders, 1959. Oocyst from Lophuromys s. stkapusi (from Levine, Bray, Ivens, and Gunders, 1959). X 2600. Fic. 260. Klossia perplexens Levine, Ivens, and Kruidenier, 1955. Oocyst from Peromyscus maniculatus (from Levine, Ivens, and Kruidenier, 1955). x 1700. Fic. 261. Isospora mcdowelli Saxe, Levine, and Ivens, 1960. Oocyst from Microtus pennsylvanicus (from Saxe, Levine, and Ivens, 1960). « 2600. Fic. 262. Isospora batabatica Musaev and Veisoy, 1960. Oocyst from Arvicola terrestris (from Musaevy and Veisov, 1960). 2100. Fic. 263. Caryospora microti Saxe, Levine, and Ivens, 1960. Oocyst from Microtus pennsylvanicus (from Saxe, Levine, and Ivens, 1960). « 2600. 315 316 PLATE 32 Fic. 264. Caryospora microti Saxe, Levine, and Ivens, 1960. Oocyst from Microtus pennsylvanicus (from Saxe, Levine, and Ivens, 1960). 2600. Fic. 265. Tyzzeria peromysci Levine and Ivens, 1960. Oocyst from Peromyscus maniculatus (from Levine and Ivens, 1960). * 2600. Fic. 266. Isospora citelli Levine, Ivens, and Kruidenier, 1957. Oocyst from Spermophilus variegatus (from Levine, Ivens, and Kruidenier, 1957). « 2600. Fic. 267. Isospora ratti n. sp. Oocyst from Rattus norvegicus (original). < 2600. Fic. 268. Dorisiella arizonensis Levine, Ivens, and Kruidenier, 1955. Oocyst from Neotoma lepida (from Levine, Ivens, and Kruidenier, 1955). x 3400. Fic. 269. Isospora ordubadica Musaev and Veisov, 1960. Oocyst from Meriones persicus (from Musaevy and Veisov, 1960). 2000. ee, ~ ea 318 PLATE 33 Fics. 270 ANp 271. Eimeria sciturorum Galli-Valerio, 1922. 270. Oocyst from “Eichhornchen” (from Yakimoff and Gousseff, 1935). X 1700. 271. Oocyst from Sciurus vulgaris (from Pellérdy, 1954). 2000. Fic. 272. Eimeria mira Pellérdy, 1954. Oocyst from Sciurus vulgaris (from Pellérdy, 1954). « 1900. Fics. 273-275. Eimeria andrewsi Yakimoff and Gousseff, 1935. 273, 274. Oocysts from ‘‘Eichhornchen” (from Yakimoff and Gousseff, 1935). x 2200. 275. Oocyst from Sciurus vulgaris (from Pellérdy, 1954). x 2000. Fic. 276. Eimeria silvana Pellérdy, 1954. Oocyst from Sciurus vulgaris (from Pellérdy, 1954). « 1900. Fic. 277. Eimeria beecheyi Henry, 1932. Oocyst from Spermophilus beecheyt (from Henry, 1932). « 2000. Fic. 278. Eimeria monacis Fish, 1930. Oocyst from Marmota monax (from Crouch and Becker, 1931). « 1900. 519 320 PLATE 34 Fic. 279. Eimeria os Crouch and Becker, 1931. Oocyst from Marmota monax (from Crouch and Becker, 1931). < 1900. Fic. 280. Eimeria perforoides Crouch and Becker, 1931. Oocyst from Marmota monax (from Crouch and Becker, 1931). x 1900. Fics. 281 aNp 282. Eimeria callospermophili Henry, 1932 from Spermophilus lateralis (from Henry, 1932). « 1800. 281. Unsporulated oocyst showing large polar granule and roughened wall. 282. Sporulated oocyst showing residual body above two sporocysts; other sporocysts not in focus. Fics. 283 anp 284. Eimeria volgensis Sassuchin and Rauschenbach, 1932. Oocysts from Spermophilus pygmaeus (from Zolotarev, 1938). * 1500. Fics. 285 anp 286. Eimeria bilamellata Henry, 1932. Oocysts from Spermophi- lus lateralis (from Henry, 1932). « 1800. 322 PLATE 35 Fic. 287. Eimeria bilamellata Henry, 1932. Oocyst from Spermophilus lateralis (from Henry, 1932). « 1800. The outer, roughened wall is split in half and lies above and below the oocyst. The inner wall is smooth and transparent. Fic. 288. Eimeria residua Henry, 1932. Oocyst from Neotoma fuscipes (from Henry, 1932). « 1800. Fics. 289 anp 290. Eimeria ondatrazibethicae Martin, 1930 emend. from Ondatra zibethica (from Martin, 1930). 289. Sporulated oocyst. & 1700. 290. Sporocyst. < 1800. Fic. 291. Eimeria citelli Kartchner and Becker, 1930. Oocyst from Spermophi- lus pygmaeus (from Yakimoft and Sokoloff, 1934). x 3600. 323 PLATE 36 Fic. 292. Eimeria beckeri Yakimoff and Sokoloff, 1935. Oocyst from Spermo- philus pygmaeus (from Yakimoff and Sokoloff, 1935). x 3600. Fic. 293. Eimeria ellobii Svanbaev, 1956. Oocyst from Ellobius talpinus (from Svanbaev, 1956). > 900. Fic. 294. Eimeria markovi Svanbaey, 1956. Oocyst from Meriones tamariscinus (from Svanbaev, 1956). x 900. Fics. 295-297. Eimeria oryzomysi Carini, 1937, from Oryzomys sp. (from Carini, 1937). 295 and 296. Sporulated oocysts. & 1300. 297. Villus, showing endogenous stages. PLATE 37 Fic. 298. Eimeria hindlet Yakimoft and Gousseff, 1938. Oocyst from Mus musculus (from Yakimoff and Gousseff, 1938). « 2200. Fic. 299. Eimeria musculi Yakimoff and Gousseff, 1938. Oocyst from Mus musculus (from Yakimoff and Gousseff, 1938). « 1700. Fic. 300. Eimeria vinckei Rodhain, 1954. Oocyst from Thamnomys surdaster surdaster (from Rodhain, 1954). & 1300. Fic. 301. Eimeria arvicanthis van den Berghe and Chardome, 1956. Oocyst from Arvicanthis abyssinicus rubescens (from van den Berghe and Chardome, 1956). & 2800. Fics. 302-312. Eimeria miyairii Ohira, 1912 from Rattus norvegicus and Rattus rattus (?) (from Matubayasi, 1938). « 3900. 302-304. Schizogony. 305. Merozoites. 306-308. Microgametogony. 309-312. Macrogametogony. 327 PLATE 38 Fics. 313-325. Eimeria nieschulzi Dieben, 1924 from Rattus norvegicus and Rattus rattus (?) (from Matubayasi, 1938). « 3900. 313-316. First type of schizogony. 317-319. Second type of schizogony. 320. Merozoites of both types. 321-323. Macrogametogony. 324 and 325. Microgametocytes. Fics. 326-332. Eimeria separata Becker and Hall, 1931 from Rattus norvegicus and Rattus rattus (?) (from Matubayasi, 1938). > 3900. 326-328. Schizogony. 329. Merozoites. 330 and 331. Macrogametogony. 332. Mature microgametocyte. Fics. 333 ANp 334. Eimeria hasei Yakimoft and Gousseff, 1936. Oocyst from Rattus rattus (from Yakimoft and Gousseff, 1936). « 1800. 529 330 PLATE 39 Fic. 335. Eimeria nochti Yakimoft and Gousseff, 1936. Oocyst from Rattus rattus (from Yakimoff and Gousseff, 1936). * 2100. Fic. 336. Eimeria ratti Yakimoft and Gousseff, 1936. Oocyst from Rattus ratius (from Yakimoff and Gousseff, 1936). & 1500. Fics. 337-339. Eimeria falciformis (Eimer, 1870) Schneider, 1875 from Mus musculus. 337, 338. Oocysts (from Wenyon, 1926). « 1500. 339. Oocyst (from Clarke, 1895). x 900. Fic. 340. Eimeria aguti Carini, 1935. Oocyst from Dasyprocta aguti (from Carini, 1935). « 1200. Fic. 341. Eimeria gliris Musaev and Veisov, 1961. Oocyst from Glis glis (from Musaev and Veisov, 1961). 2000. Fic. 342. Eimeria capibarae Carini, 1937. Oocyst from Hydrochoerus hydro- choerus (from Carini, 1937). x 1700. Fic. 343. Eimeria hydrochoeri Carini, 1937. Oocyst from Hydrochoerus hydro- choerus (from Carini, 1937). « 1700. Fic. 344. Eimeria cotiae Carini, 1935. Oocyst from Dasyprocta aguti (from Carini, 1935). « 1100. Fic. 345. Eimeria paraensis Carini, 1935. Oocyst from Dasyprocta aguti (from Carini, 1935). « 1100. 331 332 PLATE 40 Fic. 346. Eimeria schueffneri Yakimoff and Gousseff, 1938. Oocyst from Mus musculus (from Yakimoff and Gousseff, 1938). x 2500. Fic. 347. Eimeria ketlint Yakimoft and Gousseff, 1938. Oocyst from Mus musculus (from Yakimoff and Gousseff, 1938). « 1800. Fic. 348. Eimeria laviert Yakimoff and Gousseff, 1936. Oocyst from Allactaga major (from Yakimoff and Gousseff, 1936). « 2700. Fic. 349. Eimeria joyeuxt Yakimoff and Gousseff, 1936. Oocyst from Allactaga major (from Yakimoff and Gousseff, 1936). 1900. Fics. 350-357. Eimeria caviae Sheather, 1924 from Cavia porcellus. 350, 351. Sporulating and sporulated oocysts respectively (from Sheather, 1924). & 1000. 352-357. Schizogony (from Lapage, 1940). 352. First division of the nucleus of the schizont. Karyosome appears to be dividing into two unequal portions. & 2250. 353. Young schizont showing two karyosomes, formed by first division of nucleus, beginning to elongate for second division. 2250. 354. Young schizont with three nuclei showing that division of the nuclei is not synchronous. ‘Two of the nuclei show only the karyosomes and the halo of karyolymph, the extrakaryosomatic areas of these nuclei having been decolorized by differentiation of the iron hematoxylin. The extra- karyosomatic area of the third nucleus still retains some of its stain. & 2250. 355. Young schizont showing simulation of division of its nucleus. & 2250. 356. Schizont with three nuclei in the resting phase. & 2250. 357. Schizont with nuclei showing amitotic divisions. & 2250. : es 4 4 “a ot 24 - = > F : La tile ae & 333 334 PLATE 41 Fics. 358-371. Eimeria caviae Sheather, 1924 from Cavia porcellus (from Lapage, 1940). 358. Schizont or microgametocyte. The two are indistinguishable until more nuclei are present. & 1500. 359. A bundle of merozoites. Residual body absent. Nuclei in typical resting phase. « 2250. 360, 361. A bundle of merozoites. Residual body present in each. Nuclei overstained so that typical structure is obscured. & 1500. 362. Microgametocyte. & 1500. 363. Microgametocyte at stage where nuclei stain lightly. & 1500. 364. Microgametocyte showing chromatin massing along one border of the nucleus before each nucleus forms the body of a microgamete. & 1500. 365. Young microgametes being formed on surface of microgametocyte. « 1500. 366. Mature microgametes on surface of microgametocyte. & 1500. 367. Young macrogamete with nucleus in typical resting phase and a cluster of small, spherical granules. & 1500. 368. An almost mature macrogamete with a single nucleus in resting phase and its cytoplasm full of granules. & 1500. 369. A young oocyst with its outer wall only, formed from the outermost row of granules; the inner ones, which stain yellow with azan, are left. « 1500. 370. An oocyst with both its walls. A few of the granules which stain yellow with azan and form the inner wall remain in the cytoplasm. The nucleus appears flattened and distorted. & 1500. 371. Mature oocyst with its walls and nucleus in typical resting phase. X 1500. PLATE 42 Fics. 372-374. Eimeria seideli Pellérdy, 1957 from Myocastor coypus (from Seidel, 1954). 372. Immature sporocyst. « 2300. 373. Mature sporocyst. & 2300. 374. Sporulated oocyst. & 1200. Fic. 375. Isospora freundi Yakimoft and Gousseff, 1935. Oocyst from Cricetus cricetus (from Yakimoff and Gousseff, 1935). « 1600. Fic. 376. Eimeria krijgsmanni Yakimoft and Gousseff, 1938. Oocyst from Mus musculus (from Yakimoff and Gousseff, 1938). x 2200. Fic. 377. Wenyonella uelensis van den Berghe, 1938. Oocyst from Funisciurus anerythrus (from van den Berghe, 1938). « 1900. Fic. 378. Wenyonella parva van den Berghe, 1938. Oocyst from Tamiscus emini (from van den Berghe, 1938). « 1900. Fic. 379. Isospora uralicae Svanbaev, 1956. Oocyst from Apodemus sylvaticus (from Svanbaev, 1956). 900. Fics. 380-382. Wenyonella hoarei Ray and Das Gupta, 1935 from Sciurus sp. (from Ray and Das Gupta, 1935). 380, 381. Sporulated oocysts. < 1400. 382. Mature sporocyst. 2100. 337 338 PLATE 43 Fics. 383-403. Cryptosporidium muris Tyzzer, 1907 from Mus musculus (from ‘Tyzzer, 1910). 383. A minute organism which has become attached to the surface of the epithelium. It has a delicate limiting membrane and an organ of attachment. The cytoplasm stains intensely blue and contains a large vacuole, probably of lipoid material. 384. A similar organism of somewhat larger size. A delicate protoplasmic process extends from the organ of attachment. 385. In this organism there are two masses of chromatin which are ap- parently about to divide into four. 386. The four masses of chromatin are somewhat irregular and each is surrounded by a mass of cytoplasm which bulges slightly from the surface of the organism. 387. The chromatin is now situated within eight fingerlike processes which project from the surface away from the organ of attachment. 388. Mature schizont about which there is no apparent limiting membrane. The eight merozoites remain in a compact mass with a small amount of residual material. 389. Free merozoites showing variation in size. 390. A minute organism with a relatively large amount of chromatin probably destined to become a microgametocyte. 391. Microgametocyte with two masses of chromatin. 392. A later stage of nuclear division. 393. Microgametocyte in which 8 masses of chromatin are in the process of dividing into 16. 394. Mature microgametocyte showing the characteristic grouping of the microgametes over the surface of the residual material away from the organ of attachment. 395. Mature microgametocyte on the surface of which there are 15 micro- gametes. 396. Small macrogamete showing nucleus and a small mass of red staining granules. 397. Macrogamete showing the outline of the iodophilic granules, which are practically unstained. 398. Group of macrogametes, one of which has a delicate process extending from the organ of attachment. All have the red-staining granules and some contain vacuoles and hyaline globules. 399. ‘Three oocysts with immature sporozoites and a residuum. The red- staining material has become fused into a homogeneous globule within the residual mass. 400. Oocyst from which the mature sporozoites have escaped. 401. Oocyst from which the sporozoites are escaping. The rod-shaped nucleus is apparent near the sharper end of the sporozoites. 402. Mature oocyst showing the characteristic arrangement of the sporo- zoites, which are bent over the centrally situated residuum and are spirally twisted. 403. ‘I'wo macrogametes showing male and female pronuclei. Fics. 404-406. Cryptosporidium parvum ‘Tyzzer, 1912 from Mus musculus (from ‘Tyzzer, 1912). « 1800. 406 404. Two minute organisms situated in the striated border of the epithe- lium of a villus. 405. Large organism with a single nucleus, probably a schizont, attached to the surface of an epithelial cell. Above this is a schizont showing 8 mero- zoites in optical cross section arranged around a centrally situated residuum. 406. Schizont with nuclei arranged at the surface away from pole of attach- ment. PLATE 44 Fics. 407-422. Cryptosporidium parvum ‘Tyzzer, 1912 from Mus musculus (from Tyzzer, 1912). « 1800. 407. Free merozoites, some of which are grouped about a mass of residual material. To the left certain organisms which have become attached are still elongate. 408. From left to right a schizont with two nuclei, two macrogametes which appear somewhat vacuolated owing to the dissolving out of the iodophilic granules; to the right of these, an uninucleated organism. 409. Macrogamete with a minute deeply stained rod or granule near its surface and between this and the nucleus a clear space, an appearance sug- gesting a stage of fertilization. 410. Mature oocyst containing four long sporozoites. 411. Immature microgametocyte. 412. Mature microgametocyte. 413, 414. Merozoites showing change in shape and structure attending their attachment to the surface of the epithelium. 415-422. Forms illustrating schizogony. The schizonts in some instances appear larger than they actually are on account of flattening. Fig. 417 shows arrangement of chromatin masses in pairs during nuclear division. Fig. 418 shows the granular structure of the chromatin, and it as well as Fig. 420 shows a delicate limiting membrane. Fig. 422 shows more or less immature merozoites. PLATE 45 Figs. 423-429. Klossiella muris Smith and Johnson, 1902 from Mus musculus (from Smith and Johnson, 1902). 423. Infected cell from a fresh kidney. It contains a sporont within a large vacuole. ‘The spherical refractile bodies in the sporont are plastin granules. x 850. 424. Double infection. The nucleus has already divided and become distri- buted near the periphery. 737. 425. Mother sporoblast, showing nuclei near periphery, containing one to four karyosomes and a mass of granular chromatin. « 1435. 426. Longitudinal section of portion of a convoluted tubule with three in- fected cells almost occluding its lumen. The two cells to the right show the attachment of the infected cell to the basement membrane by a narrow, densely granular peduncle. The increase in size of the infected cells is clearly seen. x 870. 427. Mother sporoblast, with daughter sporoblasts beginning to form. < Wee. 428. A stage a little later than the preceding. The daughter sporoblasts, each with its nucleus at its distal end, appear to bud out from a large central mass. & 1590. 429. Daughter sporoblasts nearly complete, still attached by a narrow peduncle to the central mass. & 1590. 343 ‘jj ies 0 344 PLATE 46 Fics. 430-437. Klossiella muris Smith and Johnson, 1902 from Mus musculus (from Smith and Johnson, 1902). 430. ‘Transverse section of convoluted tubule, with very large flask-shaped cell, containing Klossiella in the daughter sporoblast stage. ‘The infected cell nearly fills the lumen. It still retains its attachment to the basement mem- brane by a narrow peduncle. & 870. 431. Infected cell from fresh kidney in salt solution. It contains 14 sporo- blasts, in which the small plastin granules are conspicuous. The cell is more or less flattened by pressure of the cover glass. & 850. 432. Small brood of sporoblasts, showing division and distribution of nuclei preparatory to formation of sporozoites. « 870. 433. Brood of sporocysts still within cell membrane of the completely atrophied host cell. Six sporocysts contain sporozoites; four are solidly stained and opaque. < 870. 434. Sporocyst from fresh kidney showing sporozoites still attached to residuum. < 850. 435. More advanced sporocyst in transverse optical section. The nucleus appears in four sporozoites. & 850. 436. Section of glomerulus showing parasite im situ in visceral layer of Bowman’s capsule. & 485. 437. Falciform bodies, oldest stage of glomerular parasite observed. & 1590. 345 346 PLATE 47 Fics. 438-443. Eimeria nieschulzi Dieben, 1924 from Rattus norvegicus (from Dieben, 1924). « 2300. 438. Unsporulated oocyst. After about half an hour the protoplasm of the new oocyst has contracted into a globule, the sporont. 439. Possible chromosome conjugation after fertilization. After a little more than one day, a clear line appears in the sporont; it extends lengthwise through the whole sporont. 440. Possible reduction division. Very often a little body appears to leave the sporont. 441. Sporont becomes irregular, and bright granules appear at four corners. ‘The sporont then becomes rectangular. 442. “Buckel” or “stringed” stage. 443. Four daughter globules formed by deepening of the grooves. 347 348 PLATE 48 Fics. 444-449. Eimeria nieschulzi Dieben, 1924 from Rattus norvegicus (from Dieben, 1924). x 2300. 444. “Pseudopyramid” stage of sporont. 445. Pyramid stage. 446. Four rounded sporoblasts formed. 447. Sporoblasts become oval and sporozoites begin to form. 448, 449. Completely sporulated oocysts. 349 350 ADDENDUM Since the manuscript for this monograph was submitted for publica- tion, several additional species of rodent coccidia have been described. Svanbaev (1963) described four species of Ezmeria and one of Isospora from the marmot Marmota marmota menzbieri (syn., M. menzbiert) in Kazakhstan, USSR. Under the name Eimeria monacis Fish, 1930, he described a species with ovoid, short ovoid, or spherical oocysts measuring 16-28 x 13-24 u, with a mean of 23.5 x 19 u; with a smooth, yellow- green, double-contoured wall 0.8-1.8 u thick; without a micropyle but with an oocyst polar granule and oocyst residuum; with spherical or ovoid sporocysts measuring 6-10.5 x 7.0—7.2 u [sic], with a mean of 8.5 x 6.5 u [sic], without a sporocyst residuum but with a refractile globule in the sporozoites. This species is neither &. monacis nor any other species of Eimeria previously described from Marmota. We are therefore naming it Eimeria tyanshanensis n. sp., since the hosts came from western Tyan- Shan. Under the name Eimeria os Crouch and Becker, 1931, Svanbaev (1963) described a species with spherical, short ovoid, or ovoid oocysts measuring 18-25 «x 16-23 u, with a mean of 22 x 20 u; with a smooth, double-con- toured, greenish or yellow-green wall 0.9-1.8 u thick; without a micropyle or oocyst polar granule but with an oocyst residuum; with ovoid sporo- cysts measuring 7-10 x 5-8 u, with a mean of 8 x 7 uw and with a sporo- cyst residuum. This species is not E. os, but appears to be E. monacis. Under the name Eimeria menzbiert Svanbaev, 1963, he described a species with ovoid or short ovoid oocysts measuring 18-28 x 13-24 u, ADDENDUM 351 with a mean of 24 x 21 u; with a smooth, double-contoured, yellow- brown or yellow-orange wall 1.2-2.1 u thick (illustrated as composed of two layers); with a micropyle but without an oocyst polar granule or oocyst residuum; with ovoid or short ovoid sporocysts measuring 8-11 < 6-8 u, with a mean of 9 x 7 u, with a Stieda body, sporocyst residuum, and refractile globules in the sporozoites. This species is probably valid, since it appears to be different from any other species of Eimeria from Marmota even though it has some resemblance to E. os. Under the name Eimeria sp., Svanbaev (1963) described a species with ovoid to short ovoid oocysts measuring 79-89 x 63-68 u, with a mean of 84 x 66 uw; with a yellow-brown, tri-contoured wall 5-6 uw thick of which the outer layer was rough and the inner layer radially striated; without a micropyle, oocyst polar granule, or oocyst residuum; with ovoid, short ovoid, or spherical sporocysts measuring 24-30 x 21-27 u, with a mean of 27 x 24 u. These oocysts did not sporulate completely. This form differs from any species of Eimeria described from Marmota, but, since it did not sporulate completely, it is best left unnamed. Under the name /sospora sp., Svanbaev (1963) described a species with short ovoid to spherical oocysts measuring 20-22 « 19-20 u, with a mean of 21 x 20 uw; with a smooth, greenish, double-contoured wall 1 u thick; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid sporocysts measuring 14-15 x 7-8 u, with a mean of 14 x 7 u. These oocysts did not sporulate completely. This form resembles /. citelli Levine, Ivens, and Kruidenier, 1957, but, since it did not sporulate completely, it is best left unnamed. Vetterling (1964) described three species of Ezmeria from the prairie dog Cynomys 1. ludovicianus from Colorado. The first was an emended description of FE. cynomysis Andrews, 1928. This species has subspherical to ovoid oocysts measuring 30-37 & 25-31 u, with a mean of 32.5 x 29 u and with length-width ratios of 1.05-1.31, with a mean of 1.15; with a smooth, yellow-green wall 1.4 u thick composed of two layers, with a mi- cropyle 5 uw wide, but without oocyst polar granule or oocyst residuum; with lemon-shaped sporocysts measuring 18 x 12 u, with a Stieda body and sporocyst residuum, without sporozoite globules. Eimeria ludoviciani Vetterling, 1964 has subspherical to ellipsoidal oocysts measuring 16-26 x 13-21 u, with a mean of 21 x 18 u and length-width ratios of 1.07-1.42, with a mean of 1.21; with a smooth, colorless wall 0.9 u thick composed of two layers; without a micropyle but with oocyst polar granule and oocyst residuum; with lemon-shaped sporocysts measuring 9 X 7 u, with a Stieda body and sporocyst residuum but without sporozoite globules. Eimeria larimerensis Vetterling, 1964 has spherical to ellipsoidal 352 THE COCCIDIAN PARASITES OF RODENTS oocysts measuring 31-40 x 26-37 u, with a mean of 36.5 x 32 u and length-width ratios of 1.00—1.33, with a mean of 1.15; with a mammillated wall composed of two layers, of which the outer is brown, granular, and 1.7 uw thick and the inner thin and colorless; without a micropyle or oocyst residuum but with an oocyst polar granule; with lemon-shaped sporocysts measuring 18 x 12 u, with a Stieda body and _ sporocyst residuum. Levine and Ivens (1963) described Eimeria siniffi Levine and Ivens, 1963 from the deermouse Peromyscus maniculatus. It has oocysts with straight sides and rounded ends measuring 26-31 x 17-19 u, with a mean of 28 x 18 u; with a smooth, colorless to pale yellowish wall com- posed of a single layer 1.2 u thick; with an operculated micropyle, with an oocyst polar granule, and without an oocyst residuum but with a trace of cobwebby material; with elongate ovoid sporocysts measuring 13-15 x 6-7 u, with a mean of 14 x 7 u, without a Stieda body but with a sporocyst residuum. Prasad (1960) described three new species of Eimeria and one of Iso- spora from different hosts. Eimeria egypti Prasad, 1960 from Meriones s. shawi has ovoid oocysts measuring 27-29 x 15-17 u, with a mean of 28 < 16 u; with a smooth, light brown wall composed of two layers of which the outer is thicker than the inner; with a micropyle 4 » wide and an oocyst residuum, but probably without an oocyst polar granule; with piriform sporocysts measuring 7-9 x 4-5 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Eimeria sylvatica Prasad, 1960 from Apodemus sylvaticus has ovoid or spherical oocysts measuring 16-21 x 10-17 u, with a mean of 18.5 x 14 u (with the spherical oocysts having a mean of 14 uw); with a smooth, pale yellow wall composed of three layers, of which the inner is very thin and the middle one rather thicker; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid sporocysts meas- uring 11-13 x 5-7 u, with a sporocyst residuum and probably without a Stieda body. Eimeria cricetomyst Prasad, 1960 from Cricetomys gambianus has ovoid or ellipsoidal oocysts measuring 18-21 x 15-17 u, with a mean of 19 x 16 uw; with a smooth, yellow wall composed of two layers of which the outer is thicker than the inner; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid sporocysts measuring 11-13 « 6-8 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Isospora egypti Prasad, 1960 from Meriones s. shawi has subspherical oocysts measuring 20-22 x 16-20 u, with a mean of 21 x 18 u; with a smooth, light brown wall composed of two layers of which the outer 1S thicker than the inner; without micropyle, oocyst polar granule, or oocyst iS) iS) 3 iS) ADDENDUM residuum; with ovoid sporocysts measuring 10-12 x 6-8 u, with a small Stieda body, with a sporocyst residuum, and with sporozoite globules. Musaev and Veisov (1962) described three new species of Eimeria from the golden hamster Mesocricetus auratus (syn., Cricetus auratus) from Transcaucasia. Eimeria aurata Musaev and Veisov, 1962 has spherical, colorless oocysts measuring 10-18 uy, with a mean of 16 u; with a smooth, yellow wall composed of a single layer 1 u thick; without a micropyle or oocyst residuum but with an oocyst polar granule; with spherical or ovoid sporocysts, the former measuring 6-8 x 4-6 u, with a mean ol 6 x 4 u, and the latter 4-6 up, with a mean of 4 u, without a Stieda body or sporozoite globules but with a sporocyst residuum. Eimeria amburdariana Musaev and Veisov, 1962 has spherical, colorless oocysts 18-24 uw in diameter, with a mean of 21 u; with a smooth wall composed of a single layer 1.5 uw thick; without a micropyle but with oocyst polar granule and residuum; with ovoid sporocysts measuring 8-12 x 6-9 u, with a mean of 11 x 8 u, without a Stieda body but with a sporocyst residuum. Eimeria razgovica Musaev and Veisov, 1962 has ovoid oocysts measur- ing 16-22 x 14-18 u, with a mean of 21 x 17 u; with a smooth wall composed of two layers each | u thick, the inner layer dark brown and the outer bright yellow; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid sporocysts measuring 6-8 x 4-6 u, with a mean of 7 x 5 u, without a Stieda body but with a sporocyst residuum and with sporozoite globules. Veisov (1962) described four new species of Eimeria from the vole Microtus majori in Azerbaidzhan. Eimeria majorici Veisov, 1962 has ovoid or ellipsoidal colorless oocysts measuring 16-28 « 12-24 u, with a mean of 23 x 17 uw; with a smooth wall composed of one layer 1.5—2.0 u thick; without micropyle, oocyst polar granule, or oocyst residuum; with piriform or ovoid sporocysts measuring 6-12 x 4-8 u, with a mean of 9 x 7 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Eimeria abuschevi Veisov, 1962 has ovoid, colorless oocysts measuring 22-31 x 16-27 u, with a mean of 29 x 25 u; with a smooth wall composed of a single layer 2 u thick; without micropyle or oocyst polar granule but with an oocyst residuum; with piriform or ovoid sporocysts 8-13 x 4-9 u, with a mean of 11 x 7 u, with a Stieda body and sporocyst residuum but without sporozoite globules. Eimeria correptionis Veisov, 1962 has ellipsoidal or ovoid oocysts measuring 18-26 x 14-20 u, with a mean of 21 x 17 u; with a smooth, yellowish wall composed of a single layer 1.5 uw thick; with a micropyle but without oocyst polar granule or oocyst residuum; with ovoid or piriform sporocysts measuring 6-11 x 4-7 uw, with a mean of 9 x 5 u, 354 THE COCCIDIAN PARASITES OF RODENTS apparently without a Stieda body, with a sporocyst residuum, and without sporozoite globules. Eimeria bicrustae Vetsov, 1962 has ovoid or ellipsoidal oocysts measur- ing 16-28 x 12-22 u, with a mean of 21 x 15 u; witha smooth wall com- posed of two layers each | u thick, the inner one dark brownish and the outer one yellowish; without micropyle, oocyst polar granule, or oocyst residuum; with ovoid or piriform sporocysts measuring 6-10 x 4-6 u, with a mean of 9 x 5 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Veisov (1963) described eight new species of Eimeria plus a ninth form to which he gave a new name from the vole Microtus arvalis in Azerbat- dzhan. He gave the new name to a form of Eimeria with ovoid, ellip- soidal, or spherical oocysts measuring 12-24 x 8-18 u, with a mean of 20 x 16 u; with a smooth, colorless to yellowish wall composed of a single layer 1.5 u thick; without a micropyle, oocyst polar granule, or oocyst residuum; with ovoid, piriform, or rarely spherical sporocysts measuring 6-10 x 4-8 u, with a mean of 9 x 7 u (and the spherical ones 4-10 u in diameter, with a mean of 6 u), with a Stieda body, sporocyst residuum, and sporozoite globules. He called it Eimeria arvalis (Iwanoft-Gobzem, 1935) nom. nov. Iwanoff-Gobzem (1935) had described it and had thought that it was the same as E. arvicolae, which Galli-Valerio (1905) had de- scribed from Microtus nivalis. Veisov (1963) said that the species of Eimeria in M. arvalis are different from those in M. nivalis, so that the present form must have a different name from FE. arvicolae. He therefore called it E. arvalis. However, the name F. arvalis is not available since, as used by Iwanoff-Gobzem (1934), it was a lapsus calami for E. arvicolae. Futhermore, Veisov’s (1963) description of this form does not differ from his own (1962) description of E. majorici from M. majori (to which he did not refer in his later paper). Hence, we consider E. arvalis of Veisov, 1963 to be a synonym of EF. majorici Veisov, 1962. Eimeria derenica Veisov, 1963 has ovoid or ellipsoidal, colorless oocysts measuring 20-38 x 14-30 u, with a mean of 29 x 21 u; with a smooth wall composed of a single layer 2-3 u thick; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid or ellipsoidal sporocysts measuring 8-16 x 4—12 u, with a mean of 12.5 x 9 u, without a Stieda body but with sporocyst residuum and sporozoite globules. Eimeria gomurchaica Veisov, 1963 has ovoid or ellipsoidal, colorless oocysts measuring 20-34 x 18-30 u, with a mean of 28.5 x 24 u; with a smooth wall composed of a single layer 2—3 u thick; without a micropyle or oocyst polar granule but with an oocyst residuum; with ovoid or piriform sporocysts measuring 8-16 x 6-12 u, with a mean of 12.5 x 9 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Or ADDENDUM 35 Eimeria zuvandica Veisov, 1963 has ovoid, rarely ellipsoidal, colorless oocysts measuring 16-26 x 12-22 u, with a mean of 22 x 16.5 u; with a smooth wall composed of a single layer 1.0-1.5 u thick; with a micropyle and oocyst polar granule but without an oocyst residuum; with ovoid sporocysts measuring 6-10 x 4-8 u, with a mean of 9 x 7 u, without a Stieda body but with a sporocyst residuum and sporozoite globules Eimeria ivadiensis Veisov, 1963 has ovoid, colorless oocysts measuring 20-32 x 16-24 u, with a mean of 25 x 20.5 u; with a rough, granulated, dark yellow wall composed of a single layer 2 u thick; without a micropyle, oocyst polar granule, or oocyst residuum; with ovoid sporocysts measuring 8-12 x 4-8 u, with a mean of 11 x 7 u, with a Stieda body and sporocyst residuum but without sporozoite globules. Eimeria monocrustae Veisov, 1963 has ovoid or subspherical oocysts measuring 22-32 x 18-28 u, with a mean of 27 x 24 u; with a rough, yellowish brown wall composed of a single layer 2.5 u thick; without a micropyle or oocyst polar granule but with an oocyst residuum; with ovoid sporocysts measuring 8-16 x 6-10 u, with a mean of 12 x 8 u, with a Stieda body, sporocyst residuum, and sporozoite globules. Eimeria twanoffi Veisov, 1963 has ovoid or ellipsoidal oocysts measur- ing 12-30 x 8-26 u, with a mean of 21.5 x 17.5 u; with a smooth wall composed of two layers each 1.25 u thick, the outer yellow-brown and the inner colorless; without a micropyle or oocyst residuum but with an oocyst polar granule; with ovoid or spherical sporocysts, the former measuring 6-14 x 4-10 u, with a mean of 10 x 7 u, and the latter meas- uring 4-8 u, with a mean of 6 u, without a Stieda body or sporozoite but with a sporocyst residuum. Eimeria primbelica Vesiov, 1963 has ovoid oocysts measuring 20-40 x 16-34 u, with a mean of 24 « 23 u; with a smooth wall 2-3 u thick com- posed of two layers, the outer yellowish brown and the inner colorless; without a micropyle but with an oocyst polar granule and oocyst resid- uum; with ovoid sporocysts measuring 6-18 x 4-12 u, with a mean of 11 x 8 u, without a Stieda body but with sporocyst residuum and sporo- zoite globules. Eimeria kolanica Veisov, 1963 has ovoid oocysts measuring 16-26 x 14-22 u, with a mean of 21 x 19 u; with a wall composed of two layers each | u thick, the outer dark brown and the inner colorless; with a micropyle but without oocyst polar granule or oocyst residuum; with spherical sporocysts 6-10 uw in diameter, with a mean of 8 u, without a Stieda body or sporozoite globules but with a sporocyst residuum. Musaev, Veisov, and Alieva (1963) described five new species of Eimeria from the social vole Microtus socialis in Azerbaidzhan. Eimeria chudatica Musaev, Veisov, and Alieva, 1963 has ovoid or ellipsoidal, colorless 356 THE COCCIDIAN PARASITES OF RODENTS oocysts measuring 10-23 x 8-21 u, with a mean of 21 x 17 u; with a smooth wall composed of a single layer 1.0—1.5 u thick; without a micro- pyle or oocyst residuum but with an oocyst polar granule; with ovoid or rarely spherical sporocysts, the former measuring 6-11 x 4—9 u, with a mean of 9 x 5 u, and the latter measuring 4-9 u, with a mean of 7 u, without a Stieda body but with a sporocyst residuum and _ sporozoite globules. This species is probably valid, although in most respects it resembles E. derenica Veisov, 1963 from M. arvalis. Eimeria cusarica Musaev, Veisov, and Alieva, 1963 has ovoid, colorless oocysts measuring 24-31 x 16-28 u, with a mean of 29 x 23 u; with a smooth wall composed of a single layer 2.0—-2.5 u thick; without a micro- pyle or oocyst residuum but with an oocyst polar granule; with ovoid sporocysts measuring 8-13 x 6-11 u, with a mean of Tl x "9s with a: Stieda body and sporocyst residuum but without sporozoite globules. This species is probably valid, although it resembles E. wenrichi Saxe, Levine, and Ivens, 1960, differing essentially only in size. Eimeria cubinica Musaev, Veisov, and Alieva, 1963 has ovoid to ellip- soidal, colorless oocysts measuring 14-24 x 10-18 u, with a mean of 21 x 15 u; with a smooth wall composed of a single layer 1.5—2.0 u thick; with a micropyle but without oocyst polar granule or oocyst residuum; with ovoid or piriform sporocysts measuring 6-11 x 4—9 u, with a mean of 9 x 5 u, with a Stieda body, sporocyst residuum, and sporozoite glob- ules. Eimeria hadrutica Musaev, Veisov, and Alieva, 1963 has ovoid or ellipsoidal oocysts measuring 16-32 x 14-24 u, with a mean of 21 x 17 u; with a smooth wall composed of two layers each 1.0-1.5 wu thick, the outer one yellowish brown and the inner one colorless; without a micro- pyle, oocyst polar granule, or oocyst residuum; with ovoid, rarely spherical sporocysts, the former measuring 6-14 x 4-12 u, with a mean of 9 x 7 u, and the latter measuring 6—9 u, with a mean of 7 u, without a Stieda body but with a sporocyst residuum and sporozoite globules. Eimeria micropiliana Musaev, Veisov, and Alieva, 1963 has ovoid or subspherical oocysts measuring 20-25 x 16-21 u, with a mean of 23 x 19 u; with a smooth wall composed of two layers, the outer one dark yellow and 1.5 u thick, the inner one colorless and 1.0 u thick; with a micropyle but without oocyst polar granule or oocyst residuum; with ovoid sporo- cysts measuring 6-11 x 4-7 u, with a mean of 10 x 5.5 u, without a Stieda body or sporozoite globules but with a sporocyst residuum. The above new species bring to 225 the number of named species of coccidia reported from rodents. Of these, 204 are Eimeria, 10 Isospora, 3 Wenyonella, 2 each Cryptosporidium and Klossiella, and 1 each Dorisiella, Caryospora, Tyzzeria, and Klossia. ~I ADDENDUM 35 LITERATURE CITED Levine, N. D., and Virginia Ivens. 1963. Eimeria siniffi n. sp. and E. arizonensis (Protozoa: Eimeriidae) from deermice in British Columbia. J. Parasitol. 49:660-661. Musaey, M. A., and A. M. Veisov. 1962. Novye vidy koktsidii roda Eimeria iz zakavkazskogo khomyaka Cricetus auratus Water (1939). Dokl. Akad. Nauk Azerbaid. SSR. 18(10):65-69. Musaev, M. A., A. M. Veisov, and F. K. Alieva. 1963. Pyat’ novykh vidov koktsidii roda Eimeria iz obschestvennoi polevki (Microtus socialis Pall.). Zool. Zhurn. 42:809-813. Prasad, H. 1960. Studies on the coccidia of some rodents of the families Muridae, Dipodidae and Cricetidae. Ann. Trop. Med. Parasitol. 54:321-330. Svanbaev, S. K. 1963. Koktsidii surka menzbira (Marmota menzbieri Kaschkarov, 1925). Akad. Nauk Kazakh. SSR. Trudy Inst. Zool. 19:51-54. Veisov, A. M. 1962. Novye vidy koktsidii roda Eimeria iz maloaziiskoi kustar- nikovoi polevki Microtus majori Thomas (1906). Dokl. Akad. Nauk Azerbaid. SSR. 18(9):59-64. Veisov, A. M. 1963. Koktsidii obyknovennot polevki Microtus arvalis Pallas (1773) v Azerbaidzhane. Izvest. Akad. Nauk Azerbaid. SSR, Ser. Biol. Med. Nauk, Baky-Baku. No. 4:61-76. Vetterling, J. M. 1964. Coccidia (Eimeria) from the prairie dog, Cynomys ludovicianus ludovicianus, in northern Colorado. J. Protozool. 11:89-91. ae - , - ‘og ’ = P a ~C ya — ie, : ¢ ad 4 ww 4 i 4) ‘ ’ 2 un 5* ‘ — © —s a aT _ ¢ sia - - ue m P89 ini : ir iF ae. & aX 7 a va Wl): ge gee iImsMtvar pagent Gare: 4, 7 : ’ : Wii ‘ap AF Cor Wire t)\.3, roelteh.o sin te a hy , LA > ow. igo J © Pre <0)? % P ‘ 4 ar Ate ; 4 ‘ ie ’ 4 * "Oh Ahan d a Rae Lng a7) i * ) ‘ _ ' we a ~ ‘ ox : + r) 5 f mee i ; 5 a aah ay ry ’ » " b, v a 7 » eee a t rea h 7 « - : Ves INDEX Acomys, 186 Adeleidae (Family), 2 Adeleorina (Suborder), 2, 3 Aguti aguti. See Dasyprocta aguti Akodon, 186 Allactaga elator, 151, 235 jaculus. See A. major major, 151, 152, 235, 332 Alticola strelzovi, 75 Anomaluridae (Family), 248 Aplodontiidae (Family), 248 Apodemus flavicollis, 107, 108, 109, 113, 114, 115, 219, 226, 227, 228, 250, 251, 296, 298 sylvaticus, 84, 106, 107, 109, 110, 111, 112, 113, 114, 115, 170, 177, 186, 226, 227, 228, 243, 296, 298, 336,-352 Arctomys marmota. See Marmota marmota Arvicanthis abyssinicus rubescens, 117, 229, 326 Arvicola arvalis. See Microtus arvalis nivalis. See Microtus nivalis terrestris, 78, 79, 80, 169, 218, 243, 286, 288, 314 Baiomys musculus, 64, 214 taylori, 64, 214, 282 Bathyergidae (Family), 164, 241, 248, 249 Bathyergoidea (Superfamily), 164, 241 Callosciurus, 186 Callospermophilus chrysodeirus. See Spermophilus lateralis Canis familiaris, 251 Caryospora, 3, 174, 186, 245 mucrot, Way 245, 31453116 Castor canadensis, 55, 211 fiber, 55 Castoridae (Family), 54, 211, 248, 249 Castorinae (Subfamily), 54, 211 Castoroidea (Superfamily), 54, 211 Cavia aperea, 153, 184, 236, 247 porcelius mos, 184,290, 24), 291, 54, 332, 334 Caviidae (Family), 152, 236, 248, 249 Caviinae (Subfamily), 152, 2 Cavioidea (Superfamily), 152, 236, 237, 238 Chinchillidae (Family), 248 Citellus. See Spermophilus Clethrionomys glareolus, 83, 84, 115, V7 TZ Oe 228,250,025 288: 290 Coccidiasina (Subclass), 2 Coccidium. See Eimeria arvicolae. See Eimeria arvicolae falciforme. See Eimeria falciformis klossiella. See Klossiella muris Coelogenus paca. See Cuniculus paca Coendou, 186 Cotia vermelha. See Dasyprocta aguti Cricetinae (Subfamily), 55, 212, 215, 248, 249 Cricetini (Tribe), 70 Cricetomys dissimilis, 147, 232, 312 gambianus, 352 Gnicetulus MmigraloniuUss\, 25 1a, 215, 216, 284, 286 Cricetus cricetus, 70, 166, 215, 242, 250, 251, 336 Cryptomys, 186 Cryptosporidiidae (Family), 2 360 THE COCCIDIAN PARASITES OF RODENTS Cryptosporidium, 2, 176, 186, 246 agutt, 157, 238, 330 muris, 176, 187, 246, 338 alactagae. See E. allactagae parvum, 178, 246, 338, 340 albigulae, 68, 215, 282 Ctenodactylidae (Family), 248 allactagae, 150, 235 Ctenomys, 186 amburdariana, 353 Cuniculidae (Family), 248 andrewst, 16, 18, 22, 201, 318 Cuniculinae (Subfamily), 156, 237 apionodes, 106, 226, 250, 296 Cuniculus paca, 156, 237 apodemt, 107, 227, 251, 296. See also Cyclospora, 34 E. rysavyt Cynomys ludovicianus, 30, 204, 250, arabiana, 100, 224, 292 260, 351 araxena, 95, 223, 290 arctomyst, 27, 203 Dasymys incomptus rufulus, 116, 228, arizonensis, 57, 212, 213, 274, 276 298 arusica, 73, 216, 286 Dasyprocta aguti, 156, 157, 238, 330 arvalis. See E. arvicolae; E. majorici Dasyproctidae (Family), 156, 237, 238, arvicanthis, 117, 229, 326 248, 249 arvicolae, 75, 217. See also E. bata- Dasyproctinae (Subfamily), 156, 238 batensis Dendromus, 186 ascotensts, 20, 202, 256 Dendromyinae (Subfamily), 248 asiatict, 32, 43, 206 Deomyinae (Subfamily), 248 assMensts, 34, 39.90, 221 Dicrostonyx groenlandicus richardsoni, astrachanbazarica, 95, 223, 290 14, 217; 286 aurata, 353 Dinomyidae (Family), 248 baiomysis, 64, 214, 282 Dipodidae (Family), 150, 235, 248, 249 batabatensis, 79, 218, 286 Dipodinae (Subfamily), 150 beckeri, 36, 41, 205, 324. See also E. Dipodoidea (Superfamily), 150, 235 talpini Dipodomyinae (Subfamily), 50, 210 beecheyi, 32, 204, 318. See also E. Dipodomys agilis, 51, 52, 210, 250 astatict brevinasus, 250 bicrustae, 354 deserti, 51, 52, 210, 250 bilamellata, 32, 204, 262, 320, 322 heermanni, 51, 52, 210, 250 bistratum, 99, 224, 292 merriamt, 51, 210, 250, 268 bohemica, 78, 218, 286 nitratoides, 51, 210 botelhoi, 13, 15, 18, 21, 23, 201 panamintinus, 51, 210, 250 callospermophilt, 34, 205, 264, 320 panamintinus mohavensis, 51, 250 callosphermophili. See E. callo- 268, 270 spermophili; E. assaensis phillipsi, 52, 210, 272 capibarae, 155, 237, 330 Dolichotinae (Subfamily), 154, 236 carini. See E. miyairit Dolichotis patagonum, 154, 236, 251, carolinensis, 61, 213, 276 312 caviae, 152, 236, 314, 332, 334 Dorisiella, 3, 171, 186, 244 cernae, 84, 219, 290 arizonensis, 171, 244, 316 chudatica, 355 Dryomys nitedula, 149, 150, 234, 312 citelli, 30, 204, 250, 260, 262, 322 correptionis, 353 Echimyidae (Family), 158, 239, 248, 249 cotiae, 157, 238, 330 Eimeria, 3, 6, 8, 186, 187, 248, 249, 250 couesit, 56, 212, 272 abuschevi, 353 coy pi, 160, 239 achburunica, 103, 225, 294 cricett, 70, 215 africana, 145, 232, 310 cricetomyst, 352 cricetult, 72, 216, 284 cubinica, 356 cusarica, 356 cynomysi. See E. cynomysis cynomysis, 29, 204, 250, 260, 351 dasymysis, 116, 228, 298 davisi, 69, 215, 284 delicata, 62, 213, 276 derenica, 354 dicrostonicis, 74, 217, 286 dipodomysis, 52, 210, 272 disaensis, 92, 222, 288 dolichotis, 154, 236, 251, 312 dorneyt, 46, 208 dura. See E. monacis dyromydis, 149, 234, 312 dzhahriana, 94, 223, 290 dzhulfaensis, 77, 217 egy ptt, 352 ellobu, 85, 219, 324 eremict, 60, 213, 276 erythrourica, 101, 225, 294 eubeckert. See E. bilamellata eutamiae, 43, 44, 206, 266 falciformis, 131, 230, 251, 308, 330. See also E. nieschulz; Eimeria sp. in Apodemus; Eimeria sp. in Cleth- rionomys glareolus; Eimeria sp. in Microtus arvalis falciformis var. criceti. See E. criceti perrst, 141, 231, 310 franchinit, 14, 18, 21, 23, 201 franklinit, 38, 205, 262 fulva. See E. seideli garnhami, 26, 203, 258, 260 geomydis, 47, 209, 266 glaucomydis, 18, 23, 44, 208, 260 gliris, 6, 148, 234, 330 gomurchaica, 354 hadrutica, 356 hall. See E. nieschulzi hansonorum, 140, 231, 308 harbelensis, 144, 232, 312 hasei, 128, 230, 328 heterocephali, 164, 241 hindlei, 135, 230, 326. See also E. prasadi; E. rysavyi; E. sylvatici; Eimeria (2) sp. in Apodemus syl- vaticus hoffmeisteri, 39, 205, 262 INDEX 361 hungaryensis, 108, 227, 251, 296 hydrochoert, 155, 237, 330 immodulata, 71, 216, 286 iradiensis, 355 twanoffi, 355 jardimlinica, 73, 216, 284 joyeuxt, Vols 235; 332 jurschuaensis, 97, 224, 292 kazakhstanensis, 36, 86, 220 keilini, 136, 230, 332. See also E1- meria sp. in Apodemus kniplingt, 22, 202, 258 kolanica, 355 krijgsmani. See E. krijgsmanni; E. svanbaevi krijgsmanni, 137, 230, 336 kriygsmanni. See E. krijgsmanni,; E. svanbaevi; E. peschankae kruidenieri, 146, 232, 314 langebarteli, 61, 213, 274 larimerensis, 351 lateralis, 38, 205, 264 laviert, 151, 235, 332 lemniscomysis, 118, 229, 298 lertkaensis, 92, 222, 288 leucopi, 63, 214, 250, 278, 280, 282 liberiensis, 144, 231, 310 liomysis, 53, 211, 272 lophuromysis, 143, 231, 310 ludoviciani, 351 luisieri, 14, 18, 22, 23, 201 mayorict, 353, 354 marionis. See E. merionis markovt, 88, 221, 324 marmotae, 27, 203 martunica, 104, 225, 294 menzbiert, 350 merionis, 87, 221 micropiliana, 356 microtina, 76, 217, 286 migratoria, 71, 215, 284 mira, 14, 18, 22, 23, 201, 250, 318 miyairit, 118, 186, 229, 298, 300, 326. See also E. nieschulzi moelleri, 17, 22, 202, 250 mohavenstis, 50, 186, 210, 250, 268, 270 monacis, 28, 204, 260, 318, 350. See also E. tyanshanensis monocrustae, 355 362 THE COCCIDIAN PARASITES OF RODENTS Eimeria (continued) muris, 105, 112, 226. See also E. he- terocephali; E. hungaryensis musajevi, 98, 224, 294 musculi, 138, 231, 326. See also E. arvicolae; E. russiensis; E. tama- riscint musculoidei, 142, 231, 310 mytairi. See E. miyairit myiari. See E. miyairit myocastori, 164, 240, 314 myopotami, 158, 239, 314 myoxt, 148, 234 nachitschevanica, 150, 234, 312 naye, 106, 112, 226 nehramaensis, 93, 222, 290 neosciurt, 19, 23, 202, 254, 256. See also E. ascotensis neotomae, 66, 214 nieschulzt, 3, 4, 5, 121, 186, 229, 251, 302, 304, 306, 328, 346, 348 nochti, 129, 230, 330 noraschenica, 90, 221, 288 nutriae, 163, 240, 314 ondatrazibethicae, 81, 219, 322 onychomysis, 65, 214, 282 operculata, 67, 215 oryzomyst, 55, 212, 251, 324 os, 29, 204, 320 paraensis, 156, 238, 330 parasciurorum, 17, 23, 45, 208, 264, 266 pellucida, 159, 239 penicillati, 49, 209, 268 perforoides, 29, 204, 320 perniciosa. See E. seideli perognathi, 49, 209, 268 peromysct, 57, 212, 274 peschankae, 89, 221 petauristae, 18, 22, 23, 46, 208, 250, 266 phyllotis, 65, 214, 251 DICH, Deen lei piriformis. See E. mira poljanski. See E. poljanskii poljanskii, 101, 225, 296 praomysis, 130, 230, 308 prasadt, 110, 227 primbelica, 355 putevelata, 117, 229, 298 rattt, 129, 230, 330 razgovica, 353 residua, 67, 214, 322 roudabushi, 62, 213, 276 rugosa, 108, 227, 251, 298 russtensis, 112, 227 rysavyt, 82, 219, 288 sadaraktica, 97, 223, 292 salasuzica, 91, 222, 288 schachtachtiana, 93, 222, 290 schamchorica, 102, 225, 294 schoutedent, 146, 232, 312 schubergr. See E. falciformis schueffnert, 139, 231, 332. See also E. cernae sciurt. See E. sciurorum sciurorum, 10, 17, 22, 23, 24, 45, 46, 201, 250, 254, 318. See also E. knip- lingi; E. moelleri; E. neosciuri; E. parasciurorum seideli, 161, 239, 336 separata, 125, 186, 229, 302, 306, 328 serbca. See E. serbica serbica, 17, 22, 23, 202, 254 sthapusit, 143, 231, 310 silvana, 16, 18, 22, 23, 202, 318 siniffi, 352 sprehni, 54, 211 stiedae. See E. ondatrazibethicae sumgaitica, 103, 225, 294 susliki, 40, 206 svanbaevi, 111, 227, 298 sylvatica, 352 tachyoryctis, 105, 226, 250, 296 talischaensis, 80, 218, 288 talpini, 38, 86, 220 tamariscini, 88, 221 tamuiasciuri, 24, 203, 258 tasakendica, 99, 224, 294 terrestris, 79, 218, 288 thomomysis, 48, 209, 268 toddi, 25, 203, 258 tyanshanensts, 350 ussuriensis. See E. beckeri; E. susliki vilast, 41, 207, 264 vinckei, 115, 228, 251, 326 vinogradovt, 96, 223, 292 volgensis, 35, 205, 262, 320. See also E. kazakhstanensis weisst, 66, 214, 251 wenrichi, 77, 218, 286 wisconsinensis, 42, 207, 264 zulfiaensis, 96, 223, 290 zuvandica, 355 sp. (in Apodemus flavicollis, A. syl- woticus), 113, 114, 227, 228, 298 sp. (in Clethrionomys glareolus), 84, 219 sp. (in Marmota bier), 351 sp. (in Microtus arvalis), 81, 219 sp. (in Rattus coucha), 131, 233 sp. (in Sciurus carolinensis), 24, 203, 258 sp. (in Scirus griseus griseus), 23, 202 sp. (in Sciurus vulgaris), 23, 203 sp. (in Spermophilus lateralis), 40, 206 sp. (in Tatera lobengulae), 147, 233 sp. (in Zygodontomys brevicauda), 69, 70, 215 (?) noellert, 156, 237 (2?) sp. (in Apodemus flavicollis), 85, 115, 228 (2?) sp. (in Apodemus sylvaticus), 114, 228 (2?) sp. (in Clethrionomys glareolus), 85, 115, 228 (?) sp. (in Nesokia indica huttoni), In7,, 233. (?) sp. (in Spermophilus parryit), 41, 206 Fimeriidae (Family), 2 Eimeriorina (Suborder), 2, 3 Eliomys quercinus, 148, 234 Kllobiini (Tribe), 85, 219 Ellobius talpinus, 36, 37, 85, 86, 87, 219, 220, 324 Erethizontidae (Family), 248 Eucoccidiorida (Order), 2 Fuspora (Subphylum), 2 Eutamias asiaticus, 32, 43, 206 dorsalis, 43, 44, 206, 266 Evotomys glareolus. See Clethrionomys glareolus marmota menz- Funisciurus anerythrus, 173, 244, 336 Geomyidae (Family), 47, 209, 248, 249 INDEX 363 Geomyinae (Subfamily), 47, 209 Geomyoidea (Superfamily), 47, 209, 210, 211 Geomys bursarius, 48, 209, 266 Gerbillinae (Subfamily), 87, 221, 248, 249 Gerbillus, 186 Glaucomys sabrinus macrotis, 46, 208 volans, 12, 17, 45, 208, 260, 264, 266 Glirinae (Subfamily), 147 Ghiroidea (Superfamily), 147, 234 Glis glis, 15, 148, 234, 250, 330 Globidium. See Eimeria Graphiurus, 186 Gregarina falciformis. See Eimeria fal- ciformis muris. See Eimeria falciformis Gregarinasina (Subclass), 2 Gymnuromyinae (Subfamily), 248 Hepatozoon balfouri. See Klossiella sp. in Jaculus jaculus jaculi. See Klossiella sp. in Jaculus jaculus Hesperomyini (Tribe), 55 Heterocephalus glaber, 164, 241 Heteromyidae (Family), 48, 209, 210, 211, 248, 249 Heteromyinae (Subfamily), 53, 211 Hydrochoeridae (Family), 155, 237, 248, 249 Hydrochoerinae (Subfamily), 155 Hydrochoerus capibara. See H. hydro- choerus hydrochoerus, 155, 237, 330 Hydromyinae (Subfamily), 248 Hystricomorpha (Suborder), 152, 236, 237, 238, 239, 241 Hystricidae (Family), 248 164, Isospora, 3, 8, 165, 186, 242 batabatica, 168, 243, 314 bigemina, 8 citelli, 165, 242, 316 egy pti, 352 freundi, 166, 242, 336 laguri, 166, 242 laguris. See I. laguri mcdowelli, 168, 243, 314 ordubadica, 169, 243, 316 364 THE COCCIDIAN PARASITES OF RODENTS Isospora (continued) ratti, 170, 243, 316 teres, 166, 242 uralicae, 169, 243, 336 sp. (in Marmota marmota menz- biert), 351 Jaculus jaculus, 185, 247 Jarrina luisiert. See Eimeria luisieri Klossia, 2, 179, 186, 247 caviae. See Klossiella cobayae perplexens, 179, 247, 314 Klossiella, 2, 3, 180, 186, 247 cobayae, 183, 247 muris, 180, 247, 342, 344 sp. (in Jaculus jaculus), 185, 24 Klossiellidae (Family), 2 ~I Lagurus lagurus, 166, 167, 242 Lemmini (Tribe), 74, 217 Lemniscomys striatus, 117, 118, 229, 298 Liomys irroratus, 53, 211 PUGIUST OS, O42 WN zi2 Lophiomyidae (Family), 248 Lophuromys stkapusi, 143, 144, 145, 146, 186, 231, 232, 310, 312, 314 Marmota bobak, 28, 204 marmota, 27, 28, 203 marmota menzbieri, 350 monax, 28, 29, 204, 260, 318, 320 Marmotini (Tribe), 27, 203 Meriones erythrourus. See M. libycus libycus, 102, 103, 104, 225, 294 PersiGuss sol 2759S, lOO a2 2 222. 243, 288, 316 shawt, 93, 94, 95, 222, 223, 290, 352 tamariscinus, 34, 35, 87, 88, 89, 90, 221, 242, 324 tristramt. See M. shawi unguiculatus, 88, 221 vinogradovi, 96, 97, 98, 99, 100, 101, 223, 224, 225, 290, 292, 294, 296 Mesocricetus auratus, 353 Microtinae (Subfamily), 74, 217, 248, 249 Microtini (Tribe), 75, 217 Microtus arvalis, 75, 81, 84, 217, 219, 250, 251, 354 mayjori, 353 nivalis, 75, 217, 354 ochrogaster, 9 pennsylvanicus, 78, 168, 174, 218, 243, 245, 286, 314, 316 socialis, 77, 217, 286, 355 Muridae (Family); 55) 2l25e ies 224- 226, 248, 249 Murinae (Subfamily), 105, 226, 248, Zao) Muroidea (Superfamily), 55, 212, 217, 22151226 Mus coucha, See Rattus coucha (Leggada) musculoides, 142, 231, 310 musculus, 9, 52, 133, 136, 137, 138, 139, 140, 141, 1775 179) 1825 iSGe230: 231,246, 247, 250; 2515 3087410326; 330; 332, 336, 338, 340 ;ea42emaa sylvaticus. See Apodemus sylvaticus Muscardinidae (Family), 147, 234, 248, 2a) Myocastor coypus, 55, 158, 159, 160, 162; 163, 164, 186, 239, 240531453326 Myomorpha (Suborder), 55, 212, 217, (apa\ i PAD, Vo, LLNS Myospalacinae (Subfamily), 248 Myoxus quercinus. See Eliomys quer- cinus Neosciurus carolinensis. See Sciurus carolinensis Neotoma albigula, 68, 69, 215, 282, 284 fuscipes, 67, 214, 322 lepida, 52, 171, 244, 250, 316 stephensi, 68, 215 Nesokia indica huttoni, 147 Ochotona pallasi, 138 Octodontoidea (Superfamily), 158, 239 Ondatra zibethica, 82, 219, 322 Onychomys leucogaster, 65, 214, 282 torridus, 52, 250 Oryctolagus cuniculus, 251 Oryzomys couest, 56, 212, 272 sp., 56, 212, 251, 324 Otomyinae (Subfamily), 248 Otomys, 186 Pedetidae (Family), 248 Perognathinae (Subfamily), 48, 209 Perognathus flavus, 49, 50, 209 formosus, 52, 250 intermedius, 49, 209, 268 longimembris, 52, 250 penicillatus, 49, 50, 209, 268 Peromyscus boylit, 52, 61, 213, 250, 274 californicus, 52, 250 eremicus, 60, 213, 276 leucopus, 9, 58, 59, 62, 63, 64, 175, 213, 214, 246, 250, 274, 276, 278, 280, 282 maniculatus, 9, 52, 58, 59, 62, 175, 180, 212, 213, 246, 247, 250, 274, 276, 314, 316, 352 nuttalli, 64, 250 men a2.91,.98, 59, 212, 250, 274, 276 Petaurista inornatus. See P. petaurista petaurista, 47, 208, 250, 266 Petauristinae (Subfamily), 44, 208 Pfeifferia schubergt. See Eimeria falci- formis Phyllotis amicus, 66, 214, 251 Pitymus, 186 Rattus (Mastomys) coucha, 131 (Dephomys) defua, 128, 229, 302 hawaiiensis, 10, 124, 128, 229, 302 moruecrous, 4,9, 52, 120, 124, 127, PAS 052295, 2305 243, 250; 251), 298, 300; 302, 304, 506, 316,. 326, 328, 346, 348 rattus, 120, 124, 127, 129, 130, 229, Zo0 20, 298, 302, 326, 328, 350 (Praomys) tullbergi rostratus, 130, 230, 308 Reithrodontomys, 186 Rhipidomys, 186 Rhizomyidae (Family), 248 Rhynchomyinae (Subfamily), 248 Sciuridae (Family), 10, 201, 208, 248, 249 Sciurinae (Subfamily), 10, 201 Sciurini (Tribe), 10, 201 Sciuroidea (Superfamily), 10, 201, 208 Sciuromorpha (Suborder), 10, 201, 208, 209; 210; 201 Semurus carolinensis, 12, 18, 19, 21, 24, 202,°203;, 250; 254,256, 258 griseus griseus, 23, 202 (Guerlinguetus) ingrami, 13, 201 INDEX 565 niger rufiventer, 12, 22, 202, 258 vulgaris, I V4 los 165 7. W839) 24, 186, 201, 202, 203, 250, 254, 318 vulgaris var. alpinus, 11, 14 sp:, 173,244, 336 Spalacidae (Family), 248 Spermophilus beecheyi, 32, 204, 318 citellus, 15, 31, 33, 204, 250 eversmannt, 37, 205 franklini, 9, 15, 33, 38, 204, 205, 262 lateralis, 33, 35, 39, 41, 204, 205, 206, 264, 320, 322 leucurus, 52, 250 maximus, 31, 35, 37, 40, 167, 204, 205, 206, 242 parryi, 41, 206 pygmaeus, 30, 31, 36, 37, 204, 205, 262, 320, 322, 324 spilosoma, 34, 35, 39, 205, 262, 264 tridecemlineatus, 31, 204, 250, 260 variegatus, 165, 242, 316 Sporozoa (Subphylum), 2 Tachyoryctes ruandae, 105, 226, 250, 296 Tachyoryctinae (Subfamily), 105, 226, 248, 249 Tamas striatus, 42, 207, 264 Tamiasciurini (Tribe), 24, 203 Tamuasciurus hudsonicus, 25, 26, 203, 258 Tamiscus eminti, 173, 244, 336 Tatera, 186 lobengulae, 147 Telosporasida (Class), 2 Thamnomys surdaster, 116, 228, 251, 326 Thomasomys, 186 Thomomys bottae, 48, 209, 268 Tyzzeria, 3, 174, 186, 246 peromysci, 174, 246, 316 Wenyonella, 3, 171, 186, 244 hoarei, 172, 244, 336 parva, 173, 244, 336 uelensis, 173, 244, 336 Xerini (Tribe), 26, 203 Xerus (Euxerus) erythropus, 27, 203, 258, 260 Zygodontomys brevicauda, 215 oat sy pe . ’ ag i eg x ob) oe aie Sant Th ne Sy = “or tne zi. “om Pa af 7 $44 me 3 raya Fe ical) eres = ud 4 q evs “i, 7 re , f- ay 7 , y ; , ; ‘ peoamarey th i a “4 . ‘ ts aro | 6 : y b ab roe ia 43 LAA Pats Fi ivi ar - a @ ai ‘ y - , i ‘ es , , E) x =) rs 7 t +- > ® L mH : — a aa c of 4 : ‘ iy ra bi 7 “ = ae = ; ae ‘ _ vere PD aoa. P38 base rer Eek SAT pseu ‘ / Sy a i oie yi: cays ad A PR PERE J j le ‘ ‘ j 4 , A) “) ‘ A 4 Wy - Vee 4