The Entomophthoraceae have been, in many classification schemes (see discussion of the order), the only family of the Entomophthorales or its equivalent. Fitzpatrick (1930) included the classical genera, Basidiobolus, Completoria, Conidiobolus, Massospora, and Empusa, in his treatment of the Entomophthoraceae. Waterhouse (1973) substituted Entomophthora for Empusa, and added Ancylistes, based upon Berdan’s (1938) study of the latter genus. Those studies (Humber, 1981a; Tucker, 1981; Ben-Ze’ev and Kenneth, 1982) that recognize three families, and of the aformentioned genera, only Entomophthora and Massospora would be included here by the latter authors. Recent students of the Entomophthorales (Humber, 1981a; Ben-Ze’ev and Kenneth, 1982) have followed Batko (1964a, 1964b, 1964c) in segregating several genera from the heterogenous genus Entomophthora (MacLeod and Müller-Kögler, 1970, 1973; MacLeod et al., 1976). Refinements of Batko’s proposals (Humber, 1981a; Ben-Ze’ev and Kenneth, 1982), and the work of Tucker (1981) on the nonentomogenous Entomophthorales, resulted in the inclusion of the following genera in the Entomophthoraceae: Completoria, Entomophaga, Entomophthora s.s., Erynia, Macrobiotophthora, Massospora, Neozygites, Strongwellsea, and Tarichium. Eryniopsis was added later by Humber (1984).

Sporangiolar dehiscence in most genera is active using papillar eversion, although Entomophthora spp. use a modification of the latter type called the “sporophore cannon.” The sporangia are not forcibly released in Massospora spp. (Tucker, 1981). These modes of sporangiolar dehiscence are discussed in detail by Benjamin (1979), Humber (1981a), and Ben-Ze’ev and Kenneth (1982). Little can assertained from resting spore ontogeny or types. The key to familes and the family description demonstrates that members of the Entomophthoraceae do have a nucleolus and the nucleus is small and stains well, and has heterochromatin (Tucker, 1981). Humber (1989) tranferred Completoria, Macrobiotophthora, and Neozygites to other families (Ancylistaceae, Completoriaceae, Neozygitaceae) and added Batkoa, Furia, and Zoophthora to the Entomophthoraceae. This is the largest family in Entomophthorales and it contains 12 genera ( Batkoa, Entomophaga, Entomophthora, Erynia, Eryniopsis, Furia, Massospora, Orthomyces, Pandora, Strongwellsea, Tarichium, Zoophthora ).

Entomophthoraceae Warming, 1884 (Haan. Syst. Bot., p. 53).

Obligate parasites of insects. Thallus consisting of well-defined hyphae, hyphal bodies, or naked protoplasts. Sporophores branched or unbranched. Spores uni- or bitunicate, discharged passively or actively by papillar eversion or on a jet of ejected cytoplasm. Resting spores zygospores or azygospores, intercalary or terminal, often formed in a budlike, nearly sessile or stalked outgrowth from one of two conjugating cells or hyphal bodies. Nuclei 5-12 µm in length, lacking a nucleolus but heterochromatin readily staining in aceto-orcein or Bismark Brown, usually uniseriate, remaining visible during mitosis; chromosomes distinct, vermiform.

Type genus: Entomophthora Fres. emend Remaudière & Keller.


A. No conidia produced; reproduction only by resting spores —— Tarichium

AA. Conidia formed; resting spores may or may not be formed —— B

B. Conidia not forcibly released, dissemination as a result of the breakdown of the abdomen of infected gregarius cicadas —— Massospora

BB. Conidia forcibly released; on insects other than gregarius cicadas —— C

C. Conidia multinucleate and unitunicate (outer wall of spore not separating from the inner layers when mounted in liquid media) —— D

CC. Conidia uninucleate and bitunicate (outer wall of spore separating from the inner layers when mounted in liquid media) —— G

D. Conidia globose to subglobose; often borne on a narrow extension of the conidiophore; pathogens of Hemiptera, Homoptera and other insect in moist regions —— Batkoa

DD. Conidia variously shaped and with a papilla; narrow extension of the conidiophore not formed; pathogens of various insects —— E

E. Conidia campanulate, papilla truncate; conidial discharge by “fungal cannon” —— Entomophthora

EE. Conidia not campanulate, papilla prominent; conidial discharge by papillar eversion —— F

F. Conidia usually globose to pyriform, multinucleate; conidiophores simple; resting spores formed —— Entomophaga

FF. Conidia fusoid to ovoid-cylindrical, with 4-12 nuclei; conidiophores simple or branched; resting spores unknown —— Eryniopsis

G. Conidiophores form a hymenium; conidia released through a hole in the abdomen of the host (flies) —— Strongwellsea

GG. Not as above —— H

H. Primary conidia ovoid to fusoid —— I

HH. Primary conidia elongate —— J

I. Rhizoids and cystidia 2-3 times as thick as the conidiophores —— Pandora

II. Rhizoids and cystidia no thicker than the conidiophores —— Furia

J. Rhizoids with holdfasts; cystidia lacking or formed no higher than the conidiophores —— Zoophthora

JJ. Rhizoids with distinct holdfasts or holdfasts lacking; cystidia grow well above the hymenium —— K

K. Conidiophores unbranched; cystida 2 times as thick, or more, than the diameter of the conidiophores —— Erynia

KK. Conidiophores digitately branched; cystidia slightly wider than the diameter of the conidiophores —— Orthomyces

Synopsis of Genera

BATKOA Humber, 1989 (Mycotaxon 34: 446); 5 spp. (Keller, 1987—ILLUS. as Entomophthora gigantea Keller, Entomophaga apiculata Thaxt.; Thaxter, 1888-ILLUS as Empusa apiculata, E. apiculata var. major Thaxt., E. papillata Thaxt.)

ENTOMOPHAGA Batko, 1964 (Bull. Acad. Polon. Sci., Sér. Sci. Biol. 12: 325) emend. Humber, 1989 (Mycotaxon 34: 447); 10 spp. (Thaxter, 1888—ILLUS. as Empusa grylli Fres.; Batko, 1964a; MacLeod et al., 1976; Humber, 1981a, 1984b; Keller, 1987).

ENTOMOPHTHORA Fresenius, 1856 [Bot. Zeitung (Berlin) 14: 883] emend. Remaudière & Keller, 1980 (Mycotaxon 11: 332); 10 spp. (Thaxter, 1888—ILLUS. as Empusa muscae Cohn and E. culicis A. Braun; Remaudière and Keller, 1980; Ben-Ze’ev and Kenneth, 1982; Descals and Webster, 1984—illus.; Keller, 1987, 2002; Keller et al., 1999).
= Empusa Cohn, 1855 (Nova Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 25: 301), non Empusa Lindley, 1824 (Bot. Reg., T. 825); fide Remaudière and Keller, 1980)
= Myriophyton Lebert, 1857 (Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 15: 26); fide Remaudière and Keller, 1980).
= Lamia Nowakowski, 1883 (Pamietn. Wydz. Akad. Umiejeten w. Krakowie 8: 173), non Lamia Endlicher, non Lamium L.; fide Remaudière and Keller (1980).
= Culicicola Niewland, 1916 (Amer. Midl. Naturalist 4: 378; fide Remaudière and Keller, 1980).

ERYNIA Nowakowski, 1881 (Dziem. III Zjazdu Lek. Pyzyr. Polak. Krakow Sekc. Bot. 6: 67) emend. Humber, 1989 (Mycotaxon 34: 449); 12 spp. (Thaxter, 1888—ILLUS., e.g., Empusa gracilis Thaxt., E. montana Thaxt., and E. rhizospora Thaxt.; Humber, 1981b; Humber and Ben-Ze’ev, 1981; Ben-Ze’ev and Kenneth, 1982; Li and Humber, 1984—illus.; Descals and Webster, 1984—illus.; Ben-Ze’ev, 1986a—illus.; Keller, 1991; 1993).
= Zoophthora Batko, 1964 (Bull. Acad. Polon. Sci. Sér. Sci. Biol. 12: 323; fide Humber and Ben-Ze’ev, 1981).
= Zoophthora Batko subgen. Erynia (Nowak.) Batko, 1966 (Acta Mycol. 2: 18; fide Remaudière and Keller, 1980).
= Zoophthora Batko subgen. Furia Batko, 1966 (Acta Mycol. 2: 20; fide Remaudière and Keller, 1980).
= Strongwellsea Batko & Weiser, 1965 (J. Invert. Pathol. 7: 460), fide Remaudière and Keller (1980).

ERYNIOPSIS Humber, 1984 (Mycotaxon 21: 258); 3 spp. (Thaxter, 1888—ILLUS. as Empusa caroliniana Thaxt. and E. lampyridarum Thaxt.; Humber, 1984a—illus.; Keller, 1991).

FURIA Humber, 1989 (Mycotaxon 34: 451); 12 spp. (Humber, 1981; Li and Humber, 1984; Thaxter, 1888—ILLUS. as Empusa virescens Thaxt., E. americana Thaxt., and E. montana Thaxt.).
= Erynia Nowakowski subgen. Furia (Batko) Li & Humber, 1984 (Canad. J. Bot. 62: 662).
= Zoophthora Nowakowski subgen. Furia Batko, 1966 (Acta Mycol. 2: 20; fide Li and Humber, 1984).

MASSOSPORA Peck, 1879 (31st Ann. Rept. N.Y. State Museum Nat. Hist., p. 44) emend. Soper, 1974 (Mycotaxon 1: 15); 12 spp (Soper, 1974—illus and key to spp., 1981—illus. and key to spp.).

ORTHOMYCES Steinkraus, Humber & Oliver, 1998 (J. Invert. Pathol. 72:5); 1 sp. (Steinkraus et al., 1998).

PANDORA Humber, 1989 (Mycotaxon 34: 451); 16 spp. (Humber, 1981, 1989; Remaudière and Hennebert, 1980; Thaxter, 1888—ILLUS. as Empusa dipterigena Thaxt. and E. echinospora Thaxt.).
= Erynia Nowakowski subgen. Neopandora Ben-Ze’ev and Kenneth, 1982 (Mycotaxon 14: 460).

STRONGWELLSEA Batko & Weiser, 1965 (J. Invert. Pathol. 7: 463) emend. Humber, 1976 (Mycologia 68: 1054); 2 spp. (Batko and Weiser, 1965—ILLUS.; Humber, 1976—illus. and key to spp., 1982).

TARICHIUM Cohn, 1870 (Beitr. Biol. Pflanzen 1: 69) emend. Lakon, 1915 (Z. Pflanzenkrankh. 25: 260), prov. name fide Humber. 1981a, for Entomophthora spp. with only resting spores known.](Macleod and Müller-Kögler, 1970—key to spp.; Keller, 1991).

ZOOPHTHORA Batko, 1964 (Bull. Acad. Polon. Sci. Sér. Sci. Biol. 12: 323); 13 spp. ( Thaxter, 1888—ILLUS. as Empusa occidentalis Thaxt.; Batko, 1964a, 1966; Humber, 1989)
= Erynia Nowakowski subgen. Zoophthora (Batko) Ben-Ze’ev and Kenneth, 1982 (Mycotaxon 14: 466).
= Zoophthora Batko subgen. Zoophthora, 1966 (Acta Mycol. 2: 16; fide Ben-Ze’ev and Kenneth, 1982b).
= Zoophthora Batko subgen. Pandora Batko, 1966 (Acta Mycol. 2: 18; fide Ben-Ze’ev and Kenneth, 1982b).

Updated Jan 31, 2009