Systematic arrangement of the sepatate gregarines

The arrangement of the septate gregarine genera presented below is based on Clopton (2002) updated to reflect new genera established since 2000 and higher level taxa recognized by Clopton (2009). I am sure there are ommissions: please notify me of corrections and provide PDF copies of supporting publications for the next site update. The right column is a simple heirarchical/alphabetical index of the septate genera and higher level groups to aid in navigation of the systematic.

Phylum Apicomplexa

Levine, 1970

All species are parasitic and have micropores, flattened subpellicular vesicles and an apical complex, generally consisting of one or more polar ring(s), rhoptries, micronemes, a conoid, and subpellicular microtubules, present at some stage in the life cycle. Cilia are absent and sexuality is by syngamy. All of the life cycle appears to be haploid except for a brief diplophase when zygotes are formed.

Class Class Conoidasida

Levine, 1988

All organelles of the apical complex are present, with the microtubular conoid consisting of a complete hollow, truncate cone. Both sexual and asexual reproduction generally occurs followed by sporogony which results in the formation of oocysts containing infective sporozoites. Cellular motility is accomplished by body flexion, gliding, undulation of longitudinal ridges or flagellar movement. Flagella are only found in microgametes of some taxa. Pseudopods are not usually formed but, if present, are used in feeding, not in locomotion. Species are homoxenous or heteroxenous.

Subclass Gregarinasina

Dufour, 1828

Mature gamonts are large and generally develop extracellularly. A mucron or epimerite is found in the sporozoites, meronts, merozoites and gamonts, with the mucron being formed from the conoid. Syzygy of gamonts generally occurs from which are formed a similar number of gametes in each gametocyte. Most often the gametes are similar, but in a few species anisogametes are differentiated with the male gamete sometimes having a flagellum or flagelliform process. Zygotes form oocysts within gametocysts thus the life cycle characteristically consists of gametogony and sporogony. The species are parasites of the digestive tract or body cavity of invertebrates or lower chordates and are generally homoxenous.

Order Eugregarinorida

Léger, 1900

Merogony absent; gamogony and sporogony present; typically parasites of annelids and arthropods but some species in other nonvertebrates; locomotion progressive, by gliding or undulation of longitudinal ridges, or nonprogressive.

Suborder Blastogregarinorina

Chatton & Villeneuve, 1936

Gamogony by gamonts while still attached to the intestine, gametes budding off of gamonts; anisogamy present; syzygy absent; gametocysts absent; oocysts with 10 to 16 naked sporozoites; gamont composed of a single compartment without definite protomerite and deutomerite, but with mucron; in marine polychaetes. This suborder contains a single monotypic family.


Chakravarty, 1960

Gamont composed of a single compartment, without definite protomerite and deutomerite, but with an epimerite or mucron in some species; syzygy present. This suborder contains ~489 named species constituting 72 genera. These animals lack a septum differentiating the body into compartments. This character distinguishes the aseptate gregarines from the septate gregarines.

Suborder Septatorina

Lankester, 1885

Gamont or trophozoite divided into protomerite and deutomerite by a septum; with epimerite, in invertebrates, especially arthropods. This suborder contains ~1,166 named species constituting 151 genera. The septate gregarines possess a septum differentiating the protomerite from the deutomerite. This character distinguishes the septate gregarines from the aseptate gregarines. Although this septum is usually apparent under light microscopy, it does not always appear complete under electron microscopy.

Superfamily Porosporicae

Chakravarty, 1960

Heteroxenous; two host species involved, one crustacean and the other molluscan. This superfamily contains a single family with 3 genera and 37 named species. Its members are differentiated from other septate gregarines by a digenic or two host life cycle.

Family Porosporidae

Labbé, 1899

Vegetative development in digestive tract of a decapod crustacean and sporogony in the connective tissue of lamellibranch molluscs.

Genus Porospora
Schneider, 1875

Oocysts absent, the sporozoites occurring in the host leukocytes; gymnospores develop into naked sporozoites rather than resistant oocysts in molluscan host; mature trophozoites relatively long, with a tendency to remain isolated. Two named species.

Genus Nematopsis
Schneider, 1892

Oocysts with a single sporozoite (monozoic); sporozoites in a doubled envelope; gymnospores develop into monozoic resistant oocysts in molluscan host; pre-reproductive associations present, typically with several individuals in straight or forked chains, intermittent; in reproductive associations primite and satellite may be enclosed in a common epicyte, protomerite deutomerite septum sometimes lost in satellite(s) which fuse to form a multinucleate compartment; protomerite of primite with a muscular collar. Thirty-two named species.

Genus Pachyporospora
Théodoridès, 1961

Gamonts with multiple nuclei, but without functional or vestigial septa. Three named species.

Superfamily Gregarinoidea

Chakaravarty, 1960 Emend. Clopton, 2009

(= Gregarinicae Chakaravarty, 1960)

Association presyzygial (trophic or gamontic); caudofrontal; gametocysts with hyaline epicyst, dehiscence by extrusion through spore tubes; oocysts liberated in monete chains. This superfamily contains 7 families with ~45 genera and ~561 named species.

Family Cephaloidophoridae

Kamm, 1922

Epimerite present; early development intracellular; association head to tail (caudofrontal), early, with primite different from satellite; marked anisogamy; gametocysts open by simple rupture; oocysts ovoid or spherical, with protruding equatorial ridge; no distinct epispore; in intestine of crustacea and other relatively primitive arthropods. This family contains 3 genera and 70 species, primarily intestinal parasites of decapods, amphipods, and cirripedes.

Genus Cephaloidophora
Mavrodiadi, 1908

Epimerite small, lenticular; oocysts ellipsoidal or spherical, with inconspicuous equatorial ring, expelled in chains or singly; in intestine of cirripedes, decapods, and amphipods. Sixty-two named species.

Genus Caridohabitans
Ball, 1959

Epimerite functional, transparent, crescent-shaped, concave anteriad; epicyte thick; nucleus with granules distributed irregularly along membrane, without endosome; gametocysts and oocysts unknown; in digestive tract of crustacea. Four named species.

Genus Rotundula
Goodrich, 1949

Epimerite button-like, persistant; trophozoite round; oocysts small, spherical or subspherical, with equatorial suture; in amphipods. Four named species.

Family Cephalolobidae

Théodoridès & Desportes, 1975

Mature trophozoites fixed to stomach epithelium by a differentiation from the protomerite; this protoepimerite is dilated into a sucker and forms lobes which adhere closely to the microvillosities of the stomach epithelium; association precocious, composed of a primite and one or two satellites, these last being placed side by side; in amphipod and decapod crustaceans. This family contains 2 genera and 5 species, primarily intestinal parasites of crustacea.

Genus Cephalolobus
Kruse, 1959

Development extracellular; association caudofrontal, occuring while attached to host gut, with one, two, or three satellites, satellites smaller than primite; protomerite of satellite without specialized holdfast; in gut of crustacea. Four named species.

Genus Callynthrochlamys
Frenzel, 1885

Trophozoite with tubules radiating out into the cytoplasm from the nucleus; syzygy caudofrontal; gametocyst oval; oocyst spherical, liberated in chains; in intestine of amphipods. Monotypic.

Family Uradiophoridae

Grassé, 1953

Epimerite simple and cylindrical; development extracellular; association caudofrontal (head to tail), precocious; with protomerite of satellite compressing deutomerite of primite; anisogamous; gametocyst opens by simple rupture; oocysts spherical, isolated, with fine equatorial ridge or radial processes. This family contains 5 genera and 21 species, primarily intestinal parasites of crustacea.

Genus Uradiophora
Mercier, 1912

Epimerite simple; development extracellular; satellite with posterior appendix; gametocysts ovoid; oocysts spherical or subspherical, with a fine equatorial backbone or ridge; gametocysts dehisce by simple rupture, oocysts not emitted in chains; in crustacea. Six named species.

Genus Heliospora
Goodrich, 1949

Epimerite simple; mature trophozoites elongate; oocysts roughly spherical, with equatorial ray-like processes formed by the epispore; in amphipods. Three named species.

Genus Pyxinioides
Trégouboff, 1912

Epimerite a ribbed button with 16 longitudinal furrows and a small apical cone, or a cupule with a central trunk; gametocysts dehisce by simple rupture; oocysts unknown or ellipsoidal, with a large appendage at one end, often united in packets; in barnacles. Ten named species.

Genus Nematoides
Mingazzini, 1891

Mature trophozoites vermiform; without septum; epimerite in form of a fork or pincers, on an elongate neck; in barnacles. Monotypic.

Genus Bifilida
Tuzet & Ormières, 1964

Epimerite unknown; oocysts cylindrical, with a long filment at each end, emitted in chains; in barnacles. Monotypic.

Family Gregarinidae

Labbé, 1899

Epimerite simple; early development intracellular; association caudofrontal, ordinarily early, even very precocious; anisogamy moderately marked; gametocysts with sporoducts; oocysts clearly elongate or cylindrical, symmetrical. This family contains 15 genera and 352 species, primarily intestinal parasites of insects.

Genus Gregarina
Dufour, 1828
(= Degiustia Levine, 1979 [in part])
(= Erhardovina Levine, 1985 [in part])

Epimerite conical, button-shaped, globular, or cylindrical; association precocious; oocysts doliform, navicular, or spherical; in intestine of insects. Three hundred and seventeen named species.

Genus Clitellocephalus
Clopton & Nolte, 2002

Epimerite deltoid, basal width at attachment to protomerite roughly equal to protomerite width, with an internal anterior obconoid structure and basal toroidal vacuole, retained in gamont; protomerite broadly elliptical to cylindrical; deutomerite narrowly obovate; association precocious, caudofrontal, biassociative, the satellite protomerite engulfing the posterior end of the primite deutomerite to form an interlock; gametocysts spherical, sporoducts present but reduced, irregular in number; oocysts dolioform, dehiscing in chains.

Genus Gymnospora
Moniez, 1886

Gametocyst with up to 8 spore ducts; with spherical oocysts. Monotypic.

Genus Triseptata
H. Hoshide, 1958

Trophozoites biassociative, cylindrical, with body composed of three segments separated by septa; epimerite a simple ovoid knob; oocysts ellipsoidal, extruded in chains. Monotypic.

Genus Gamocystis
Schneider, 1875

Epimerite absent; protomerite transitory, lost in mature trophozoites; association precocious; trophozoite body cylindrical, round, or tongue-shaped; oocysts cylindrical; in intestine of insects. Seven named species.

Genus Anisolobus
Vincent, 1924

Epimerite apparently absent, even in youngest stages; anterior margins of protomerite forming a strong adhesive disk; association very early, precocious; gametocysts ellipsoidal, with thick wall; 6-8 sporoducts present; oocysts doliform, emitted in chains; in insects. Four named species.

Genus Anisoloboides
Théodoridès, 1992

Similar to Anisolobus. Eugregarines with bi-associative trophozoites, with globular shape, globular shape conspicuous in primite; protomerite with 15-20 well-differentiated lobes. Gametocysts spherical; oocysts unknown. Four named species.

Genus Garnhamia
Crusz, 1957

Epimerite papillate to acicular; without septum between protomerite and deutomerite (weak hyaline septum?); association takes place while the primite is still attached to midgut epithelium of host; gametocysts with sporducts; oocysts ovoid, extruded in chains; in silverfish. Monotypic.

Genus Torogregarina
Geus, 1969

Biassociative; protomerite with a broad swelling around its base; gametocysts round; oocysts ovoid. Monotypic.

Genus Faucispora
Baudoin, 1967

Oocysts with ellipsoidal endospore and with epispore distended at both ends to form spouts closed by a small cap-valve. Monotypic.

Genus Spinispora
Baudoin, 1967

Oocysts fusiform, covered with spines over their whole surface. Monotypic.

Genus Bolivia
Corbel, 1968

Trophozoites filiform; protomerite with an anterior depression and often with bulb at base; deutomerite cylindrical, with fine longitudinal striations; gametocysts spherical, with mucilaginous sheath, with a single, short broad sporoduct and a large residuum; oocysts doliform, not emitted in chains but in an agglomerated mass. Monotypic.

Genus Cirrigregarina
Levine, 1979

Similar to Gregarina (epimerite conical, button-shaped, globular or cylindrical; oocysts doliform, navicular or spherical); found in barnacles. Two named species.

Genus Molluskocystis
Levine, 1979

Similar to Gregarina (epimerite conical, button-shaped, globular or cylindrical; oocysts doliform, navicular or spherical); found in mollusks. Monotypic.

Family Blabericolidae

Clopton, 2009

Gregarinoidea with caudofrontal gamontic association. This family contains 2 genera and 9 species, all parasites in cockroaches.

Genus Blabericola
Clopton, 2009

Epimerite simple, globular to pyriform, without diamerite; deutomerite of gamont obpanduriform; association gamontic, presyzygial, caudofrontal; gametocysts with spore tubes, dehiscing by extrusion of monete oocyst chains; oocysts elliptoid to dolioform; parasites of blaberid cockroaches.

Genus Protomagalhaensia
Pinto, 1918

Development intracellular; young trophozoites always elongate; associations caudofrontal, with toungue-like interlock; oocysts dolioform, with corner spines, released from gametocyst by simple rupture. Three named species.

Family Metameridae

Levine, 1979

Epimerite simple; trophozoite with secondary segmentation of protomerite and deutomerite; in annelids. This family contains 4 genera and 5 species, all parasites of annelids.

Genus Metamera
Duke, 1910

Epimerite subconical, apex eccentric, with many branched digitiform processes; gametocysts dehisce by simple rupture; oocysts biconical, navicular; in leeches. Two species.

Genus Gopaliella
Ganapati, Kalavati & Sundaram, 1974

Epimerite umbrella-like, with a central, deeply staining rod; body with many segments (usually 8, occasionally 11); isogamous; oocysts spherical, with 8 sporozoites; gametocysts dehisce by simple rupture. Monotypic.

Genus Deuteromera
Bhatia & Setna, 1938

Epimerite subconical, with a cup-shaped apex; protomerite and deutomerite of trophozoite with incomplete secondary segmentation; syzygy, gametocysts, and oocysts unknown; in polychaetes. Monotypic.

Genus Cognettiella
Pizl, Chalupský & Levine, 1983

Epimerite simple; protomerite and deutomerite of trophozoite with complete secondary segmentation; syzygy, gametocysts, and oocysts unknown; coelomic, in polychaetes. Monotypic.

Family Didymophyidae

Léger, 1892

Septum of satellite resorbed more or less slowly during syzygy; gametocysts spherical or somewhat elongate; oocysts with a loose epispore and an ellipsoidal endospore; oocysts emitted in packets enveloped by a thin membrane (secondary cysts) by means of a single, very long sporoduct with a thin wall. This family contains 1 genus and 41 species, all parasites of colepterans.

Genus Didymophyes
Von Stein, 1848

Epimerite cylindroconical, very degenerate, reduced to a small, pointed papilla; deutomerite with needle-like cyrstaloids of unknown chemical nature in endoplasm; gametocyst usually spherical;oocysts grouped in spherical packets in gametocysts. Forty-one named species.

Family Hirmocystidae

Grassé, 1953

Epimerite ordinarily papilla-like or simple knob-like; gametocysts dehisce by simple rupture; oocysts ellipsoidal, prismatic, fusiform, ovoid, or even spherical. This family contains 16 genera and 69 species.

Genus Hirmocystis
Labbé, 1899

Epimerite a conical button or papilla, labile; trophozoites often associated in chains; oocysts ovoid, ellipsoidal, or cylindroid, in insects. Thirty-seven named species.

Genus Neohirmocystis
Ghose, Ray, & Haldar, 1986

Epimerite apparently absent or vestigial; association caudofrontal, early (although some solitary individuals may remain); protomerite deutomerite septum present in satellite; gametocysts dehisce by simple rutpture; oocysts spherical. Two named species.

Genus Hyalospora
Schneider, 1875

Epimerite a globular button; young trophozoite endoplasm orange-yellow; oocysts ellipsoidal or fusiform. Nine named species.

Genus Tettigonospora
L. M. Smith, 1930

Similar to Hirmocystis; epimerite spherical; primite resembling satellite; gametocysts dehisce by simple rupture; oocysts hyaline and spherical. Monotypic.

Genus Naiadocystis
Clopton, Cook & Cook, 2004

Epimerite a simple cordoid or toroid epimerite with an interior obconoid structure resembling a funnel that tapers to a distinct axial canal bisecting the protomerite, enveloped by an accessory form–fitting membrane, conspicuous in all stages of development but reduced in satellites in association; satellite protomerite reduced to a linearly crateriform cup or sucker that receives and enfolds posterior end of primite deutomerite; association precocious, caudofrontal, biassociative; gametocysts spherical, hyaline coat present; sporoducts present but reduced, irregular in number; oocysts axially symmetric, broadly elliptoid with 4 small spherical polar knobs, 1 each at 308, 1508, 2108, and 3308, relative to the longitudinal axis of the oocyst, extruded en masse through rupture of sporoducts or by simple rupture in loose chains that immediately dissipate as individual oocysts in water. Four named species.

Genus Dumbbellicephalus
Bala & Kaur, 1988

Epimerite spherical with basal constriction, contriction in anterior portion of protomerite giving the appearance of a"dumbbell"; satellite with a distinct"pad", broad, anteriad, forming junction with primite; gametocysts dehisce by simple rupture, oocysts spherical or ovoid. Monotypic.

Genus Euspora
Schneider, 1875

Sporonts solitary; association caudofrontal; gametocysts dehisce by simple rupture, without sporoducts; oocysts prismatic. Four named species.

Genus Tintinospora
H. Hoshide, 1959

Trophozoites in associations of two or three; epimerite a simple globular papilla; gametocysts dehisce by simple rupture; oocysts extruded in chains. Monotypic.

Genus Arachnocystis
Levine, 1979

Epimerite simple, spherical; gametocysts spherical, dehiscing by simple rupture, without residuum; oocysts biconical, with truncate ends; in arachnids. Three species.

Genus Acanthogregarina
Kalavati, Narasimahamurti & Vnidyullatadevi, 1988

Epimerite a disc with approximately 14 peripheral indentations, arranged in two whorls of 7 indentation each; biassociative; gametocysts without sporoducts, oocysts cylindrical to dolioform with thickened poles, released in membrane-bound masses by rupture of the gametocyst. Monotypic.

Genus Eliptocystis
Sengupta, Ghosh & Haldar, 1991

Early development extracellular; young trophozoites elongate; associations caudofrontal, (with notched interlock); oocysts spherical, released from elliptical gametocyst by simple rupture. Monotypic.

Genus Pintospora
Carini, 1944

Trophozoites at first intra- and then extracellular; gametocysts with smooth wall; oocysts unite in pairs, each one with thickened, rather pointed ends, with smooth walls, frequently free in coelomic cavity of host. Monotypic.

Genus Endomycola
Théodoridès, Desportes, & Jolivet, 1972

Trophozoites solitary and globular, with button-like epimerite and spherical nucleus; entocyte and ectocyte very chromophilic; gametocysts and oocysts unknown. Two named species.

Genus Retractocephalus
Haldar & Chakraborty, 1976

Epimerite globular, retractile into protomerite; initial development intracellular; association head to tail (caudofrontal); oocysts dolioform, liberated from the gametocyst in chains by simple rupture. Six named species.

Genus Liposcelisus
Sarkar & Haldar, 1980

Epimerite spherical, hyaline, with a corona of 14 to 16 ridges; gametocysts simple, dehiscing by simple ruture; oocysts spindle-shaped. Monotypic.

Genus Quadruhyalodiscus
Kundu & Haldar, 1984

Epimerite spherical, with a corona of four anterior sucker-like hyaline disks, with short neck; trophozoites biassociative; gametocysts dehisce by simple rupture; oocysts spindle-shaped (narrowly elliptic), with polar thickenings; development extracellular. Monotypic.

Superfamily Stenophoroidea

Levine, 1984 Emend. Clopton, 2009

(= Stenophoricae, Levine, 1984)

(= Solitaricae Chakravarty, 1960)

Association syzygial; frontal or laterocaudofrontal. Gametocysts with hyaline epicyst, dehiscence by expulsion of single oocysts en masse by internal pressure of gametocyst residuum, expulsion of oocysts in polyete chains by an internal gametocyst residuum; rupture of a gametocyst valve and expulsion of oocysts in monete chains by an internal gametocyst residuum, or expulsion of unchained oocysts en masse in response to hyaline epicyst constriction or an internal gametocyst residuum. This superfamily contains 14 families with ~103 genera and ~568 named species.

Family Stenophoridae,

Léger & Duboscq, 1904

Early development intracellular; epimerite absent or reduced to an invaginated form; syzygy head to tail, late (just before encystment); anisogamy accentuated, gametocysts open by simple rupture, eliminating oocysts singly; oocysts ovoid or ellipsoidal, with very ample epispore, with or without equatorial suture; in diplopods. This family contains 3 genera and 93 species, all intestinal parasites of millipedes.

Genus Stenophora
Labbé, 1899
(= Stenocephalus Schneider,1875)
(= Spirosoma Ishii, 1915)

Development intracellular; oocysts ovoid, with prominent equatorial suture. Eighty-eight species.

Genus Fonsecaia
Pinto, 1918

Development intracellular; oocysts elongate ovoid, without equatorial ridge. Monotypic.

Genus Hyalosporina
Chakravarty, 1935

Epimerite a diskoid collar (bordered with fine, invasive rootlets in mature trophozoites), with central tongue-like papillae; without neck; nucleus attached to posterior end of deutomerite by two lateral bundles of myonemes; anisogamous; oocysts ovoid, with large polar epispore. Four species

Family Leidyanidae

Kudo, 1954

Similar to Gregarinidae, (epimerite simple; early development intracellular; syzygy caudofrontal, anisogamy moderately marked; gametocysts with sporoducts; oocysts clearly elongate or cylindrical, symmetrical), but solitary; epimerite simple, knob-like, gametocysts with several sporoducts; oocysts dolioform. This family contains a single genus, Leidyana, with 29 named species.

Genus Leidyana
Watson, 1915

Syzygy late; epimerite a simple globular knob; oocysts liberated from gametocyst through spore ducts (dehiscence by spore ducts); spores dolioform. Twenty-two species.

Family Cnemidosporidae

Grassé, 1953

Young trophozoite at first fixed to intestinal cell by a voluminous epimerite which, in the course of development, regresses and becomes reduced to a small cap or spherule on the protomerite; sporoducts absent (?); oocysts ellipsoidal, in intestines of millipedes. This family contains a single genus, Cnemidospora, with 4 named species.

Genus Cnemidospora
Schneider, 1882

With the characters of the family. Four named species.

Family Monoductidae

Ray & Chakravarty, 1933

Initial development intracellular; epimerite present or absent, if present, with prongs; gametocyst with a single sporoduct; oocyst ellipsoidal or ovoid, with hyaline epispore; octozoic (oocyst with eight sporozoites). This family contains 4 genera and 32 species, primarily intestinal parasites of millipedes.

Genus Monoductus
Ray & Chakravarty, 1933

Epimerite a small elevation, with basally attached prongs, with a deep myocyte composed of two axial bundles, one retracting the epimerite; anisogamous; oocysts ellipsoidal to fusiform, with mediodorsal ridge; released from the gametocyst in a chain through a single sporoduct, in millipedes. Five named species.

Genus Stenoductus
Ramachandran, 1976

Epimerite absent in mature trophozoite; oocysts ellipsoidal, with hyaline epispore, without mesodorsal ridge, extruded in chains; nucleus without myonene tethers of Monoductus, in millipedes. Twelve named species.

Genus Phleobum
Haldar & Chakraborty, 1974

Epimerite absent; gametocyst with ectocyst, endocyst, and single sporoduct; oocysts oval, not extruded in chains; in insects. Two named species.

Genus Pyxinia
Hammerschmidt, 1838
(= Xiphorhynchus Léger, 1892)
(= Beloides Labbé, 1899)

Epimerite a crenulate crateriform disk with or without hooks at its periphery, with a central conical style; gametocysts dehisce by formation of a hole in the wall through which oocysts are extruded in a single thread; oocysts biconical; in beetles. Thirteen named species

Family Sphaerocystidae

Chakravarty, 1960 Emend. Clopton, 2009

Epimerite simple, globular, labile; trophozoites and gamonts subspherical to spherical; limits of protomerite and deutomerite septum marked by distinct differences in cytoplasmic density and granularity, protomerite deutomerite septum ephemeral; oocysts ovoid. Monotypic. This family contains 1 genus and 4 species, primarily intestinal parasites of insects.

Genus Sphaerocystis
Léger, 1892

Epimerite labile; trophozoites apparently monocystic, solitary, parthenogenetic (?); oocysts ellipsoidal, with epispore; in insects. Four named species.

Family Trichorhynchidae

Ormières, Marquès, & Puisségur, 1977

Epimerite slightly lobed, carried on a long neck, limited by three unit membranes; epicyte ornamented by numerous digitations forming a dense"beard"; epicytic folds absent but arches and longitudinal"apical" filaments regularly disposed in the membranes; gametocyst studded with teats or nipples [Fr. mamelons]; dehiscence by means of a lateral residuum ("pseudocyst") gametocysts often with an equatorial suture; oocysts cylindrical, emitted in chains; in chilopods. This family contains 1 genus with 2 species, both intestinal parasites of centipedes.

Genus Trichorhynchus
Schneider, 1882

With the characters of the family. Two named species.

Family Dactylophoridae

Léger, 1892

Epimerite complex, digitate, without a long neck; nucleus in protomerite; syzygy (association) late, frontal (head to head); anisogamy very accentuated; gametocyst generally with a single residuum ("pseudocyst") coming from the male and serving for expulsion of the oocysts; oocysts cylindrical, with rounded ends; in intestine of chilopods and sometimes millipedes and insects. This family contains 8 genera and 29 species, primarily intestinal parasites of centipedes.

Genus Dactylophorus
Balbiani, 1899

Protomerite dilated, with a large number of"baguettes"; gametocyst spherical; oocysts in more or less long chains. Two named species.

Genus Echinomera
Labbé, 1899

Epimerite an irregular cone ending in an eccentric point, bearing a number of short digitiform rhizoids; gametocysts without sporoducts, with residuum formed by male gamont; oocysts cylindrical, in chains. Eight named species.

Genus Grebnickiella
Bhatia, 1938

Protomerite spread out transversely, with numerous delicate rhizoids, made up of two long, narrow horizontal lobes fused and turned up spirally at one end, peripheral portion with many teeth from which long filaments project; oocysts long, ovoid, mostly in chains. Nine named species.

Genus Rhopalonia
Léger, 1894

Epimerite a subspherical button, with ten or more digitiform processes; gametocyst with residuum ("pseudocyst"); oocysts cylindrical. Three named species.

Genus Acutispora
Crawley, 1903

Epimerite a button at the end of a long neck; gametocyst with or without residuum; oocyst biconical, with a thick, blunt endocystic rod at each end. Three named species.

Genus Seticephalus
Kamm, 1922

Epimerite forming a thick tuft of short, anteriorly directed, brush-like bristles; protomerite broad and flat. Monotypic.

Genus Dendrorhynchus
Keilin, 1920

Epimerite a disk or sucker with a border of ramified lobes (papillae); septate; transverse fibrils (epicytic, not forming septa) conspicuous, often giving the protodeutomerite a metameric appearance; gametocysts ellipsoidal, irregular; oocysts fusiform. Two named species.

Genus Mecistophora
Ganapati & Narasimhamurti, 1960

Epimerite cup-shaped with numerous intracellular filaments; protomerite deutomerite septum present; gametes dissimilar; gametocyst with residuum; oocysts simple, octozoic. Monotypic.

Family Amphiplatysporidae

Kundu & Haldar, 1984

Epimerite absent; solitary, association late, lateral; gametocysts ovoid with prominent ectocyst (pseudocyst), dehiscing by simple rupture; oocysts cylindrical, with polar plates, extruded in chains. This family contains a single monotypic genus, Amphiplatyspora.

Genus Amphiplatyspora
Kundu & Haldar, 1984

Epimerite absent; solitary, association late, lateral; gametocysts ovoid with prominent ectocyst (pseudocyst), dehiscing by sinple rupture; oocysts cylindrical, with polar plates, extruded in chains. Monotypic.

Superfamily Stylocephaloidea

Clopton, 2009

Association syzygial; frontal or frontolateral. Gametocysts with gelatinous or papyriform epicyst, dehiscence by simple rupture or dissolution; oocysts liberated singly or in monete chains by simple exposure.

Family Stylocephalidae

Ellis, 1912

(= Stylorhynchidae Schneider, 1875)

Epimerite elongated, with or without appendages; development extracellular; syzygy (association) frontal, late; gametocyst residuum present; oocysts purse- or hat-shaped, emitted in chains, with a single brown or blackish wall, with a dehiscence line on the convex border; in arthropods. This family contains 15 genera and 91 species, all parasites of insects.

Genus Stylocephalus
Ellis, 1912
(= Stylorhynchus Schneider, 1875)

Epimerite elongated into a neck, with subapical tumidus; gametocysts covered by papillae, with residuum ("pseudocyst"); oocysts purse-like, usually emerging in long chains in insects. Thirty-seven named species.

Genus Stylocephaloides
Théodoridès, Desportes, & Jolivet, 1965

Epimerite a button; older trophozoites with epicytic ribs; protomerite often with thin axis; oocysts purse-like; in insects. Two named species.

Genus Cystocephalus
Schneider, 1886

Epimerite with short neck, broad at base, swollen into a bulb, and surmounted with a small, olive-shaped prolongation; oocysts hat-shaped; in insects. Ten named species.

Genus Bulbocephalus
Watson, 1916

Epimerite consisting of stout, broad-based style borne on a long neck, with a central bulbous swelling; oocysts unknown; in insects. (Note: The oocysts of Bulbocephalus indicus Narain, 1961 are tear-drop shaped Narain [1961].). Three named species.

Genus Xiphocephalus
Théodoridès, 1964 Emend. Corbel, 1971

Epimerite very long, terminating in a sharp or blunt point, with a filiform neck. Eight species.

Genus Lophocephalus
Labbé, 1899
(= Lophorhynchus Schneider, 1882)

Epimerite a large cupule formed by a crenulate crown, with longitudinal striations and basal border, with or without a crown of small membraneous vesicles; nucleolus in the shape of a long, twisted sausage; oocysts hat-shaped, asymmetrical, in insects. Two named species.

Genus Lophocephaloides
Théodoridès, Desportes, & Jolivet, 1965

Young trophozoites with a long, button-shaped epimerite on a short neck; epimerite of older trophozoites and gamonts a more or less flacid floral corolla somewhat resembling that of Lophocephalus; gametocysts of the usual cystocephalid type; oocysts in strings like beads; in insects. Two named species.

Genus Sphaerorhynchus
Labbé, 1899
(= Sphaerocephalus Schneider, 1886)

Epimerite a small sphere at the end of a long or sometimes short neck; in insects. Four named species.

Genus Oocephalus
Schneider, 1886

Epimerite a rounded button borne on a short conoidal neck hardly longer than wide; in insects. Two named species.

Genus Campanacephalus
Théodoridès, 1955

Epimerite without neck, consisting of a quadrangular or truncated button; oocysts emitted in chains; in beetles. Two named species.

Genus Clavicephalus
Théodoridès, 1959

Epimerite a quadrangular knob which prolongs the protomerite, finely punctate at apex, with a finely punctate"islet" differentiating the cytoplasm of the epimerite from that of the protomerite; syzygy head to tail; gameotcyst and oocyst unknown; in insects. Three named species.

Genus Cystocephaloides
Théodoridès & Jolivet, 1963

Epimerite with a short neck, consisting of a button, surrounded by an epicytic collarette; with development the collarette disappears and the epimerite takes the form of a quadrangular button carried on a short neck; in insects. Monotypic.

Genus Orbocephalus
Nelson, 1970

Epimerite a dilated papilla surrounded by a flat disk; borne on a long neck (diamerite); gametocysts and oocysts unknown. Two named species.

Genus Lepismatophila
Adams & Travis, 1935

Epimerite at first elongated into a neck but then turning into a simple, symmetrical knob; mature trophozoites with a broad protomerite; gametocyst with a residuum; gametocyst walls ornamented with channels delimited by small granular areas; oocysts ovoid or ellipsoidal, assymentrical (hat-shaped), liberated from the gametocyst in uncoiling chains by simple rupture. Ten named species.

Genus Colepismatophila
Adams & Travis, 1935

Epimerite a simple symmetrical knob, globular; oocysts hat-shaped, with polar filaments, curved; released from the gametocyst in wavy chains. Three species.

Family Actinocephalidae

Léger, 1892

Epimerite symmetrical, with or without appendage, syzygy (association) late, frontal; gametocysts without sporoducts, generally dehiscing by simple rupture; oocysts biconical, cylindrobiconical or irregular; simple or spiny; in intestine of arthropods or chordates. This family contains 3 subfamilies, 61 genera and 283 species.

Subfamily Actinocephalinae

Léger, 1899

Oocysts without spines or thickenings at their poles; in insects, diplopods, chilopods, opilionids, and chordates. This subfamily contains 31 genera and 137 species.

Genus Actinocephalus
Von Stein, 1848
(= Stephanophora Léger, 1892)

Epimerite a hemispherical tumidus with eight to ten upwardly directed digitform processes, borne on a short but prominent neck; neck persists more or less in trophozoite, but digitiform processes disappear; gametocysts dehisce through a hole in the gametocyst wall through which oocysts are extruded in a single thread; oocysts biconical or lemon-shaped; in insects. Forty named species.

Genus Caulocephalus
Bhatia & Setna, 1924

Epimerite dilated into a caulifower shape anteriorly, narrower at base, persistant; protomerite with specialized cytoplasmic zone; oocysts ovoid or spherical in beetles. Three named species.

Genus Cornimeritus
H. Hoshide, 1959

Trophozoites solitary; epimerite an acute claw with a long, large, flexible stalk; gametocysts spherical, dehiscing by simple rupture; oocysts biconical, extruded in lateral chains. Monotypic.

Genus Umbracephalus
H. Hoshide, 1959

Trophozoites solitary, elongate cylindrical, with elongate ellipsoidal nucleus; epimerite with a very long neck bearing an anterior crown with about twenty recurved hooks; gametocysts spherical; oocysts unknown. Monotypic.

Genus Urnaepimeritus
H. Hoshide, 1959

Trophozoites solitary, elongate, with spherical nucleus; epimerite with a short neck and a bowl-shaped crown with 30 or more recurved hooks; gametocysts spherical, dehiscing by simple rupture; oocysts spindle-shaped. Monotypic.

Genus Pilidiophora
Baudoin, 1967

Epimerite without a differentiated neck and having the form of a small bonnet consisting of a dome prolonged toward the base by little tongues surrounding the diamerite. Monotypic.

Genus Geneiorhynchus
Schneider, 1875 Emend. Clopton & Cook, 2007

Trophozoites solitary, epimerite an orbicular, hemispherical or toroidal tumidus bearing many semifalciform or semiluniform spines, borne on an intercalating diamerite; association laterofrontal, late, just prior to syzygy; oocysts hesperidiform, released from gametocysts by simple rupture. Six named species.

Genus Acanthoepimeritus
H. Hoshide, 1959

Trophozoites solitary, with spherical nucleus; epimerite a swollen club, with nine or ten rows of hooks around it and with numerous recurved hooks covering its anterior surface. Monotypic.

Genus Phialoides
Labbé, 1899
(= Phialis Léger, 1892)

Epimerite with a long neck, consisting of a retractile papilla bordered by a cushion set peripherally with stout teeth, surrounded by a wider collarette; gametocysts spherical, without sporoducts; oocysts biconical, ventricose; in beetle larvae. Monotypic.

Genus Legeria
Labbé, 1899
(= Dufouria Schneider, 1875)

Epimerite with a stalked, irregulary lobed, and folded plasma portion; protomerite surrounded by a collar; gametocysts without sporoducts; oocysts cylindrobiconical or subnavicular, with a thick wall; in beetle larvae. Monotypic.

Genus Steinina
Léger & Duboscq, 1904

Epimerite a short, motile, digitiform process which later changes into a flattened button; oocysts biconical, ventricose; in insects. Twenty-three named species.

Genus Bothriopsides
Strand, 1928
(= Bothriopsis Schneider, 1875)

Epimerite without neck, lenticular, small, sessile, oval, with six or more long, slender, filaments projecting anteriad; protomerite very large, mobile; protomerite deutomerite septum convex anteriad; gametocysts spherical; oocysts biconical. Eight named species.

Genus Pomania
Baudoin, 1967

Epimerite composed of a point which may be tranformed into a sucker and a cap covering the protomerite. Monotypic.

Genus Stictospora
Léger, 1893 Emend. Hays & Clopton, 2004

Epimerite compound with terminal epimerite holdfast proper and short intercalating diamerite; holdfast very broadly obovoid, with a margin of sympetalous, pendulate, narrowly to very narrowly elliptoid lamina, variable in number, epimerite absent in mature gamonts; association late, laterofrontal, biassociative; gametocysts spherical, hyaline coat present; sporoducts absent; oocysts irregularly dolioform with distinct lateral sulcate margins and slight axial keel; released en masse by simple rupture of gametocyst. Seven named species.

Genus Coleorhynchus
Labbé, 1899
(= Coleophora Schneider, 1885)

Epimerite diskoid, poorly developed; protomerite large, forming a collarette or sucker with a muscular sucker serving for attachment; trophozoites solitary; development parthenogenetic; oocysts biconical or navicular, octozoic; in intestine of insects. Monotypic.

Genus Amphoroides
Labbé, 1899
(= Amphorella Léger, 1892)

Epimerite a globular sessile papilla, lost early; protomerite globular, cup-shaped anteriad; oocysts biconical or navicular, curved, without epispore; in millipedes. Four named species.

Genus Stylocystis
Léger, 1899

Epimerite simple, a sharply pointed process, normally recurved in a long hyaline point or spine, very sharp anteriad; protomerite absent; gametocysts mature entirely within host and do not contain residuum; oocysts biconical; in intestine of insects. Four named species.

Genus Taeniocystis
Léger, 1905

Epimerite sessile or with short neck, consisting of an apical tumidus with six to eight recurved hooks; deutomerite divided by septa into many serial segments; gametocysts spherical, opening by simple rupture; oocysts biconical; in insects. Three named species.

Genus Sciadiophora
Labbé, 1899
(= Lycosella Léger, 1896)

Epimerite a large sessile, disk with a crenulate border, lost early; protomerite with numerous vertical lamellae, broadening to an umbrella in the mature trophozoite, each rib of umbrella curved to form a spine pointing backward; oocysts biconical or ovoid, united into a string of beads; in opilionids. Six named species.

Genus Anthorhynchus
Labbé, 1899
(= Anthocephalus Schneider, 1887)

Epimerite a large flattened, fluted disk; oocysts biconical or ovoid, in lateral chains. Three named species.

Genus Agrippina
Strickland, 1912

Epimerite a circular disk, with peripheral digitiform processes; oocysts symmetrical, ellipsoidal, unarmed; in flea larvae. Monotypic.

Genus Alaspora
Obata, 1953

Epimerite sessile in a jar-shaped sucker, with a deeply depressed anterior end; trophozoite solitary, elongate; gametocyst opens by simple rupture; oocysts cylindrobiconical, with three thin, triangular, longitudinal plates extending radially from the trunk; in intestine of beetles. Monotypic.

Genus Ascocephalus
Obata, 1955

Epimerite sessile, sucker-like, with a thick peripheral collar, depressed very deeply at anterior end, with cavity widened at bottom and a rugged ring of tooth-like projections around posterior periphery; trophozoites solitary, elongate ovoid; gametocyst spherical; gametocyst dehiscence and oocysts unknown; in intestine of beetles. Monotypic.

Genus Amphorocephalus
Ellis, 1913

Epimerite with a short dilated neck, consisting of a sessile button fluted on its sides; protomerite with a transverse superficial constriction; in chilopods. Seven named species.

Genus Tricystis
Hamon, 1951

Only gamont known; intracellular or between cells of digestive epithelium; epimerite, protomerite, and deutomerite present; in chaetognaths. Two named species.

Genus Thalicola
Ormières, 1965

Trophozoites with or without longitudinal striations; epimerite simple; syzygy head to tail; gametocysts spherical; oocysts and mode of dehiscence unknown; in salps. Four named species.

Genus Epicavus
Ormières & Daumal, 1970

Epimerite in the form of a deep cup with thick walls slightly striated longitudinally on their inner surface, carried on a neck; nucleus with a single large nucleolus; gametocysts spherical, dehiscing by simple rupture; oocysts subspherical, with rounded episporal polar plugs; in insects. Two named species.

Genus Gryllotalpia
Hasselmann, 1926

Epimerite a large knob set on a long stalk; gametocysts dehisce by simple rupture; oocysts biconical; in insects. Monotypic.

Genus Chilogregarina
Levine, 1979

Epimerite simple or with nonpersistant digitiform processes at apex; gametocysts and oocysts unknown; in chilopods. Five named species.

Genus Harendraia
Sarkar, 1984

Trophozoites ovoid to fusiform; epimerite with narrow, elongated neck, consisting of a highly-complex,long, bowl-like structure with a bulb-like, round base, apex truncated, set with 4 short, slender symmetrical filaments or spines; mature trophozoites solitary, cylindrical; gameotcysts spherical, dehiscing by simple rupture; oocysts ellipsoidal, released in lateral chains. Monotypic.

Genus Levinea
Kori, 1985

Early trophozoites solitary; epimerite cup-like at the apex of a short neck, with numerous peripheral digitiform processes; dehiscence of gametocysts by simple rupture; oocysts cylindrobiconical; in odonate insects. Monotypic.

Subfamily Pileocephalinae

Baudoin, 1967

Epimerite globular; oocysts biconical, without spines or thickenings at their poles; in insects. This subfamily contains 10 genera and 46 species.

Genus Asterophora
Léger, 1892

Epimerite a thick, horizontal disk with a milled border and a stout style projecting from the center; oocysts cylindrobiconical; in intestine or arthropods. Fourteen named species.

Genus Schneideria
Léger, 1892

Epimerite a thick horizontal disk with milled border, with or without a central style; protomerite deutomerite septum lost (resorbed) early, oocysts biconical, smooth, swollen; in insects. Five named species.

Genus Paraschneideria
Nieschulz, 1924

Epimerite simple, button-like; development intracellular; mature trophozoites without protomerite deutomerite septum; oocysts biconical; in insects. Monotypic.

Genus Neoschneideria
Ormières, Manier, & Mathiez, 1965

Epimerite composed of a lobed pad in the center of which is a sharp papilla; epicyte longitudinally striated; trophozoite divided into segments, generally six; gametocyst with external wall forming two long prolongations opposite each other; oocysts biconical, with very loose epispore, emitted in chains; in insects. Two named species

Genus Neoasterophora
Cook & Clopton, 2011

Association late, caudofrontal; epimerite in the form of an apical ovoidal tumidus; diamerite present, with milled toroidal disk; oocysts fusiform in outline, without polar plugs; oocyst residua present as a central cluster; released from gametocysts by simple rupture. Monotypic.

Genus Pileocephalus
Schneider, 1875

Epimerite lance-shaped or a triangular button, with short neck; oocysts biconical; in insect larvae. Eighteen named species.

Genus Gemmicephalus
Baudoin, 1967

Epimerite in the form of an oval bud; oocysts slightly biconical. Two named species.

Genus Discorhyncus
Labbé, 1899
(= Discocephalus Léger, 1892)

Epimerite a large spheroidal papilla with collar and short neck, non-persistant; oocysts biconical, slightly curved, slightly ventricose; in insect larvae. Monotypic.

Genus Globulocephalus
Baudoin, 1965

Young trophozoite with globular epimerite with a permanent septum; syzygy late and ephemeral, intermediate between frontal and lateral; gametocysts dehisce by simple rupture; oocysts biconical; in intestine of trichopteran larvae. Monotypic.

Genus Crucocephalus
Sarkar, 1984

Young trophozoites ovoid, becoming cylindrical; epimerite with short, thick-walled neck, dilated at base, consisting of a globular tumidus with 6-8 broad, vertical, peripheral ridges; mature trophozoites fusiform to cylindroconical, solitary; gametocysts spherical, dehiscing by simple rupture; oocysts biconical, with sharply-pointed ends. Monotypic.

Subfamily Acanthosporinae

Léger, 1892

Oocysts with spines or thickenings at their poles; sometimes at the equator and also along their edges; in carnivorous insects (especially aquatic insects), chilopods, and opilionids. This subfamily contains 20 genera and 75 species.

Genus Acanthospora
Léger, 1892

Epimerite a conical papilla with an obtuse point; or simple, knob-shaped, papilla-like; or crateriform, with partial septa around it; oocysts biconical or ellipsoidal, with polar and equatorial spines, liberated from the gametocyst by simple rupture of the gametocyst wall; in insects. Six named species.

Genus Grenoblia
Hasselmann, 1927

Epimerite small, simple; protomerite conical; gametocysts and oocysts unknown. Monotypic.

Genus Corycella
Léger, 1892

Epimerite with a central button bearing a crown of strong, recurved hooks that terminate in acute tips; oocysts biconical, with a tuft of fine hairs at the ends. Two named species.

Genus Ancyrophora
Léger, 1892

Epimerite consisting of a globular papilla with flexible or rigid appendages forming hooks which may be recurved; oocysts biconical, with polar tuft and 6 equatorial bristles or spines. Twenty-two named species.

Genus Rhizionella
Baudoin, 1971

Epimerite consisting of a central button bearing long, ascending, filamentous, root-like appendages; oocysts biconical, without polar truncation, with polar knobs and equatorial spines. Monotypic.

Genus Cometoides
Labbé, 1899
(= Pogonites Léger, 1892)

Epimerite with neck or stalk, consisting of a spherical or flattened papilla surrounded by long, thin, flexible filaments; gametocysts dehisce by simple rupture; oocysts cylindroconical, with polar spines and two rows of equatorial spines in intestine of insects. Four named species.

Genus Prismatospora
Ellis, 1914

Epimerite subglobose with lateral, recurved hooks; oocysts prismatic, composed of a central, regular hexagonal prism capped at each end by a rectangular, truncated hexagonal pyramid; spines long, in two series of 6 each, inserted symmetrically in the tetrahedral angles at the junctions of the apical pyramids with the central prism; in dragonfly naiads. Monotypic.

Genus Nubenocephalus
Clopton, Percival & Janovy, 1993

Epimerite broadly ovoid, truncated posteriad, with broad, flexible equatorial tumidi that do not form hooks, spines, or digitiform processes; borne on a long, slender stalk; oocysts dodecahedral, elongate, terminally truncate, hexagonal in equatorial cross section, without equatorial faces, with equatorial and terminal spines. Monotypic.

Genus Tetraedrospora
Tschudovskaia, 1928

Epimerite consisting of a flattened disk, bordered by 14 to 16 hooks; gametocysts develop and oocysts emerge in host gut; cysts elongate tetrahedral, with sides bearing a row of spines; in fungus gnats. Monotypic.

Genus Ramicephalus
Obata, 1953

Epimerite dishlike, with many upward-projecting dentritic processes around the periphery; oocysts biconical, with one row of polar and one row of six equatorial spines. Thirteen named species.

Genus Calyxocephalus
Clopton, 2004

Holdfast compound, comprising a terminal epimerite and intercalating diamerite, epimerite a thick disk or linearly crateriform sucker with distal apopetalus calyx of petaloid lobes, diamerite short (less than half of the total holdfast length) very broadly obdeltoid; association late, cephalolateral, biassociative; gametocysts spherical, hyaline coat present, conspicuous sporoducts absent, dehiscing by simple rupture to release individual oocysts en masse; oocysts axially symmetric, hexagonal dipyramidic in shape with slight polar truncations, bearing 6 equatorial spines, 1 at each equatorial vertex, 6 terminal spines obliquely inserted at each pole, 1 at each vertex created by polar truncation. Monotypic.

Genus Quadruspinospora
Sarkar & Chakravarty, 1969

Epimerite spherical or subspherical, with variable number of stumpy, digitiform processes directed laterally; gametocysts thick-walled, spherical, dehiscing by simple rupture; oocysts ovoid, with two very long spines at each pole, gamonts, solitary, in insects. Seven named species.

Genus Coronoepimeritus
H. Hoshide, 1959

Trophozoites solitary; Epimerite with a short neck, consisting of a crown-like, globular tumidus, covered with many small digitiform processes which may or may not be branched; oocysts ellipsoidal or ovoid, with long, filament-like polar spines, released from the gametocyst by simple rupture. Nine named species.

Genus Dinematospora
Tuzet & Ormières, 1954

Development extracellular; epimerite hemispherical, flattened, becoming button-shaped with trophozoite maturity, attached by a short, broad, stalk to the protomerite at a chromophilic ring that persists until syzygy; deutomerite with a thick membrane; longitudinal myonemes well developed; paraglycogen granules and black granules present; nucleus spherical or slightly ovoid, with a single nucleolus; young trophozoites solitary; gametocysts dehisce by simple rupture; oocysts with 2 long polar filaments. Monotypic.

Genus Doliospora
Ormières & Baudoin, 1969

Epimerite without ornamentation; oocysts asymmetrical, without terminal tufts, with two equatorial thickenings on the longitudinal cordons; in opilionids. Two named species.

Genus Acanthosporidium
Georgévitch, 1951

Middle part of anterior end of epimerite in the form of a tapered snout; with a short neck separating the epimerite and protomerite; gametocysts and oocysts unknown. Two named species.

Genus Cosmetophilus
Cockendolpher, 1991

Locomotion progressive by gliding and undulation. Gametocysts passed in the host's feces without sporoducts, dehiscing by simple rupture. Oocysts biconical with thickenings at poles and equator with row of spines on poles and along edges, released by the thousands but not in chains. Trophozoite divided into protomerite and deutomerite by septum. Epimerite symmetrical without hooks, filaments, striations, or ornamentation. Trophozoites, sporonts, and gametocysts develop in the intestine and intestinal cecae of cosmetid Opiliones. Monotypic.

Genus Contospora
Devdhar & Amoji, 1978

Epimerite without neck, consisting of a conical tumidus with basal dentition and approximately 20 vertical lamellae; oocysts cylindrical, tapering to blunt polar points, with tufts of spines at each pole; in arthropods. Monotypic.

Genus Tetractinospora
Sarkar & Haldar, 1981

Epimerite a globular holdfast with more than 8 (~ 16) laminate, vertical (retroarcate) plates; borne on a short neck; trophozoites solitary; gametocysts dehisce by simple rupture; oocysts biconical, bent in the middle, with 2 sharp, stout spines at each pole. Monotypic.

Genus Echinoocysta
Levine, 1984

Epimerite a simple globular or spherical knob; protomerite dome-shaped or hemispherical, with striated rim around its base, set on a short cylindrical collar; oocysts biconical, with a row of 8-10 slender spines at each end; oocysts released from gametocyst in chains of 2 to 3 or more by simple rupture. Monotypic.

Genus Mukundaella
Sarkar, 1981

Epimerite consisting of a very short neck ending in a cup with numerous striations or vertical folds; oocysts diamond-shaped, hexagonal in polar view, with polar and meridional spines; in insects. Three named species.

Genus Tetrameridionospinospora
Kori & Amoji, 1985

Epimerite consisting of a short but distinct neck ending in a globular or discoidal bulb with numerous peripheral tentacles or digitiform processes; oocysts biconical with four polar spines (two at each pole) and four meridional spines (two on each side). Two named species.

Subfamily Menosporinae

Léger, 1892

Oocysts crescentic, smooth; in arthropods. This subfamily contains 5 genera and 24 species.

Genus Menospora
Léger, 1892

Epimerite very persistent, consisting of a long neck terminated by a cupule bordered by hooks; oocysts bent, banana-shaped (crescentic); in insects. Two named species.

Genus Hoplorhynchus
Carus, 1863

Epimerite consisting of a long neck terminated by a flattened disk bordered by hooks or spines; oocysts crescentic or ellipsoidal and curved. Twelve named species.

Genus Odonaticola
Sakar & Haldar, 1981

Epimerite consisting of a long neck terminated by an umbrella shaped bulb with marginal petaloid spines; trophozoites solitary; gametocysts dehisce by simple rupture; development extracellular; oocysts smooth, navicular. Eight named species.

Genus Domadracunculus
Clopton, 1995

Epimerite borne on a narrow basal stalk; a pleated cup or sucker, very broadly ovoid in lateral view, circular en face; with distinct pleats rising from the interior of the cup to form a rosette or crown. Gametocysts spherical; hyaline coat erratic (not present on all gametocysts); sporulation by simple dehiscence; no spore tubes or packet membranes were observed. Oocysts crescentic. Development solitary; association lateral, late. Monotypic.

Genus Steganorhynchus
Percival, Clopton & Janovy, 1995

Epimerite set on a long vermicular stalk; an ovoid papilla enclosed in a retractile, globular sheath. Gametocysts spherical; hyaline coat present; sporulation by simple dehiscence; no spore tubes or packet membranes were observed. Oocysts crescentic. Development solitary; association lateral, late. Monotypic.

Family Brustiophoridae

Kundu & Haldar, 1981

Trophozoites solitary; epimerite small, variously formed, with fine bristles, with short but distinct neck; gametocysts dehisce by simple rupture; oocysts spherical, set with brush borders and chained with fine filamentous processes. This family contains 1 monotypic genus described from the intestine of a beetle.

Genus Brustiospora
Kundu & Haldar, 1981

Epimerite consisting of a short but distinct neck ending in a bundle of fine bristles; gametocysts dehisce by simple rupture; oocysts spherical, set with brush borders and connected by fine, filamentous processes into a chain. Monotypic.

Family Acutidae

Stejskal, 1965

Trophozoites solitary; epimerite simple, changing shape during development; gametocysts without sporoducts or residuum, dehiscing by opening in gametocyst wall; oocysts ellipsoidal, in insects. This family contains 2 montypic genera, both described from the intestines of domestic honey bees.

Genus Acuta
Stejskal, 1965

Epimerite simple, growing continually, at first spherical, then cylindrical and drop-shaped and breaking off, leaving a scar; gametocysts dehisce by rupture of a simple, irregular fissure in the gametocyst wall; in intestine of bees. Monotypic.

Genus Apigregarina
Stejskal, 1965

Epimerite simple, at first large, becoming ovoid, then spherical, and finally conoidal in the course of development; gametocysts dehisce by rupture of a round fissure in the gametocyst wall; in intestine of bees. Monotypic.

Family Monoicidae

Geus, 1969

Autogamy present (i.e. each gamont forms gametes of both sexes). This family contains 1 monotypic genus described from the intestines of domestic honey bees.

Genus Monoica
Stejskal, 1964

Epimerite simple; young trophozoites solitary; each gamont forms gametes of both sexes; gametes anisogamous; gametocysts dehisce by simple rupture; gametocyst residuum present; oocysts ellipsoidal; in bees. Monotypic.

Superfamily Fusionicae

Stejskal, 1965

Homoxenous; upon syzygy the nucleus and entocyte of the satellite go into the primite, where they fuse; anisogamous; gametocysts and oocysts unknown. This superfamily is monotypic.

Family Fusionidae

Stejskal, 1965

With the characters of the superfamily.

Genus Fusiona
Stejskal, 1965

With the characters of the family, in insects. Monotypic.

Heirarchical/alphabetical index

Suborder Septatorina
Superfamily Fusionicae
Family Actinocephalidae
Subfamily Acanthosporinae
Genus Acanthoepimeritus