Mucoraceae
Overview
The concept of Mucoraceae has undergone a number of changes in the last few years from the one presented as part of a comprehensive survey of Mucorales by Hesseltine and Ellis in 1973, which refected the thinking of most students of the Mucorales up to that time and on to the end of the decade (Benjamin, 1979). As treated by Hesseltine and Ellis (1973) Mucoraceae included some 19 to 21 genera of Mucorales having only more or less large, multispored, columellate sporangia, although in a few genera (e.g., Backusella, Pirella, Sporodiniella, Syzygites) traditionally placed in the family, some or all of the sporangia are greatly reduced in size or otherwise modified. Subsequent to the review by Hesseltine and Ellis (1973) several genera were added to the Mucoraceae; these were described as new [ Halteromyces (Shipton and Schipper, 1975); Thermomucor (Subrahamanyam et al., 1977; Apophysomyces (Misra, Srivastava, and Lata, 1979], Circinomucor (von Arx, 1982), and Micromucor von Arx, 1982 [formerly a subgenus of Mortierella; Gams, 1977]), reinstated ( Mycocladus [Mirza et al., 1979]), or realigned ( Umbelopsis (Amos and Barnett, 1966 [described originally as a hyphomycete] but now in the Umbelopsidaceae). Several realignments of genera restricted the limits of Mucoraceae by removing a number of genera to other families. Benny and Benjamin (1975, 1976) placed Pirella and, following Pidoplichko and Mil’ko (1971), Backusella in the Thamnidiaceae because of their terminal deliquescent sporangia and lateral, persistent-walled sporangiola. Mirza (Mirza et al., 1979) established the new family, Dicranophoraceae, for species of Dicranophora, Spinellus, Sporodiniella, and Syzygites. Finally, von Arx (1982) further limited the circumscription of Mucoraceae by establishing the Absidiaceae for those genera formerly included in the family which have a more or less apophysate sporangium; von Arx also created the Phycomycetaceae for Phycomyces and Spinellus. Umbelopsis is now included in the Umbelopsidaceae.
Absidiaceae is considered a synonym of Mucoraceae and Dicranophoraceae is treated as a synonym of Thamnidiaceae by Hawksworth et al. (1995) and Kirk et al. (2001). All of the remaining families of the Mucorales are being treated as synonyms of Mucoraceae based on the phylogenetic study of the Mucorales published by O’Donnell et al. (2001).
There are references to the history of the Mucorales in the discussion for each of the families that have been palced in synonomy with the Mucoraceae. The reader has access to the discussion of these families in the following manner.
Mucoraceae are common saprobes in soil, dung, and other decaying organic material.
Classification
Mucoraceae Dumortier 1822 ( Com. Bot., p. 69 et 82).
= Pilobolaceae Corda ex Corda, 1842 ( Anleit. Stud. Mycol., p. 71).
? Pilobolaceae Corda, 1837 ( Icon. Fung. 1: 22).
= Chaetocladiaceae Schröter, 1893 (In Engler and Prantl’s, Die Nat. Pflanzenfam. 1(1): 131. 1893 emend. R.K. Benjamin & Benny, 1993 (Mycologia 85:670).
= Choanephoraceae Schröter 1893 [In Engler and Prantl, Die natürl. Pflanzenfam. 1(1): 131]
= Thamnidiaceae Fitzpatrick, 1930 ( The Lower Fungi Phycomycetes, p. 253).
= Syncephalastraceae Naumov ex Benjamin, 1959 (Aliso 4: 327).
≡ Syncephalastraceae Naumov, 1935 Opredelitel Mukorovykh ( Mucorales ), Ed. 2, p. 16 ( nomen nudum, without a Latin diagnosis; Art. 36.1 of the ICBN, Greuter et al., 2000)].
= Cunninghamellaceae Naumov ex R. K. Benjamin, 1959 (Aliso 4: 415).
≡ Cunninghamellaceae Naumov, 1935 [ Opredelitel Mukorovykh ( Mucorales ), Ed. 2, p. 16; nomen nudum, without a Latin diagnosis, Art. 36.1 of the ICBN (Greuter et al., 2000)].
= Radiomycetaceae Hesseltine & Ellis, 1974 (In Ellis and Hesseltine, Mycologia 66: 91).
≡ Radiomycetaceae Hesseltine & Ellis, 1973 [In Ainsworth, Sparrow, and Sussman. The Fungi 4B, p. 202 nomen nudum, without a Latin diagnosis; Art. 36.1 of the ICBN (Greuter et al., 2000)].
= Saksenaeaceae Hesseltine & Ellis, 1974 (In Ellis and Hesseltine, Mycologia 66: 87).
≡ Saksenaeaceae Hesseltine & Ellis, 1973 (Ainsworth, Sparrow, and Sussman (Eds.). The Fungi 4B, p. 202 ( nomen nudum, without a Latin diagnosis; Art. 36.1 of the ICBN, Greuter et al., 2000)].
= Dicranophoraceae Mirza, 1979 (In: Mirza et al., Mucorales of Pakistan, p. 19).
= Absidiaceae v. Arx, 1984 [1982] (Sydowia 35: 21).
= Phycomycetaceae v. Arx, 1984 [1982] (Sydowia 35: 22).
= Mycotyphaceae Benny & R. K. Benjamin, 1985 (In Benny, Kirk, and Samson, Mycotaxon 22: 122)(Benny et al., 1985—KEY TO GENERA).
= Gilbertellaceae Benny, 1991 (Mycologia 83: 151).
Saprobic or a facultative gall-forming parasites of other Mucorales. Mycelium well developed; rhizoids and stolons often present. Sporangiophores erect or ascending, simple or branched, arising from substrate mycelium, aerial hyphae, or stolons and rhizoids. Sporangia similar, more or less globose, columellate or acolumellate, uni- or multispored, with or without an apophysis; wall diffluent, deliquescent, persistent, or persistent with preformed zones of rupture. Suspensors formed on aerial hyphae or sporangiophores, opposed, apposed, or tong-like, equal or unequal, often swollen, with or without appendages, may be wrapped around one another. Zygospores with a pigmented, usually dark, thick-walled zygosporangium that is ornamented with warts, spines, or ridges, it can be more or less translucent and undulate to smooth; smooth-walled, translucent zygosporangia enclosing a pigmented zygospore can also be formed; suspensors opposed, apposed, or “tongs-like,” more or less equal or heterogamous, usually unadorned or may bear appendages.
Type genus: Mucor Micheli ex L.: Fr.
KEY TO THE GENERA OF MUCORACEAE
A. Sporangia flasked-shaped with an elongate, slender neck deliquescing at the tip, multispored—— Saksenaea
AA. Sporangia otherwise, globose to obpiriform, rarely another shape, uni- to multispored——B
B. Sporangiospores formed mostly in a single row in merosporangia borne on a vesicle—— Syncephalastrum
BB. Not with the above combination of characteristics——D
C. Sporangiospores often broadly fusiform-elliptical, with a striate wall (sometimes faint), often with apical, hyaline appendages; sporangial wall persistent, with a longitudinal suture——D
CC. Not with the above combination of characteristics——E
D. Sporulates well after repeated transfer on most laboratory culture media; sporangiola not produced; zygospores with rough walls, suspensors opposed; commonly causes soft rot of peaches—— Gilbertella
DD. Sporulates well after repeated transfer only on nutritionally weak media (e.g., potato-carrot agar) or on susceptable host plants; sporangia only or sporangia and/or sporangiola produced; zygospores with striate walls, suspensors apposed; saprobes or weakly parasitic on susceptable flowering plants——KEY 1
E. Large columellate sporangia never produced; pedicellate, uni- or multispored sporangiola developed on small or large vesicles——F
EE. Large columellate sporangia produced; these sometimes accompanied by sporangiola——J
F. Primary vesicles on sporophore give rise to simple, uniseptate branches, which form terminal, secondary vesicles bearing uni- or multispored, pedicellate, acolumellate sporangiola——KEY 2
FF. Not with the above combination of characteristics——G
G. Sporangiola columellate, uni- or multispored; saprobic or gall-forming parasites; sterile spinelike branches sometimes formed——H
GG. Sporangiola acolumellate, uni- or multispored; sterile spinelike branches sometimes produced; some taxa parasitic on other fungi——I
H. Sporangiola unispored; sterile spines may be formed; one genus a gall-forming parasite of other Mucorales in nature, readily grows in the laboratory on ordinary culture media——KEY 3
HH. Sporangiola uni- or multispored, sterile spines not produced; saprobic——KEY 4 (Couplets G-O; in part, Cokeromyces, Ellisomyces, Kirkomyces, Phascolomyces, Zychaea, and sometimes Pirella, Thamnidium, and some spp. of Thamnostylum )
I. Sporangiola uni- or multispored; sporangiospores brownish, wall striate; zygospores with apposed suspensors——KEY 1 (in part, Choanephora, Blakeslea )
II. Sporangiola unispored; sporangiospores hyaline, smooth- or spinose-walled; zygospores with opposed suspensors——J
J. Colonies characteristically low growing, nonmucoroid, dark brown or gray; sporophore simple, or rarely once or twice branched; fertile vesicles terminal; sporangiola uni- or multispored, cylindrical, globose, or obovoid, only smooth; sporangiola and pedicels mono- or dimorphic; sporangiolar wall persistent; sporangiolar pedicel dehisces by a basal preformed circumscissile zone of rupture——KEY 5
JJ. Colonies characteristically cream colored to light yellow or gray, mucoroid; sporophores mostly branched, fertile vesicles terminal and/or lateral; sporangiola unispored, globose to elliptical, usually spinose; sporangiola and pedicels monomorphic; sporangiolar wall deliquescent; sporangiolar pedicels dehisce by random breakage—— Cunninghamella
K. Deliquescent-walled sporangia and persistent-walled sporangiola developed together——KEY 4 (Couplets B-F ; in part, Backusella, Pirella, Thamnidium, Helicostylum, Fennellomyces, Thamnostylum )
KK. Only sporangia developed, these with fugacious or deliquescent walls; sporangiola not formed——L
L. Sporangia more or less apophysate——M
LL. Sporangia nonapophysate; sporophore may be vesiculate or constricted, especially immediately below the sporangium——N
M. Mostly saprobic, one sp. of Absidia a facultative parasite of other Mucorales——KEY 6
MM. Parasitic in nature on mushrooms or insects——KEY 7
N. Sporangial wall dark, persistent; subsporangial vesicle present or absent; sporangium forcibly discharged or passively released as a unit upon contact with substratum——KEY 8
NN. Not with the above combination of characteristics——O
O. Sporophore unbranched, up to 5 cm or more long, usually blue-green, with metallic sheen; sporangium up to 500 μm in diam; heterothallic, zygospores developed between tonglike, appendaged suspensors—— Phycomyces
OO. Sporophore branched or unbranched, rarely up to 5 cm in length, usually colorless to yellowish or grayish, sporangia usually less than 200 μm in diam (but see Mucor plasmaticus van Tiegh. [Schipper, 1975]); hetero- or homothallic; zygospores developed between opposed, nonappendaged suspensors——KEY 9
KEY 1
A. Only relatively large, columellate sporangia produced—— Poitrasia
AA. Sporangiola always present, usually accompanied by sporangia——B
B. Sporangiole dehiscent; wall separable, with longitudinal line or lines of weakness—— Blakeslea
BB. Sporangiole indehiscent; wall not separable, without longitudinal lines of weakness—— Choanephora
KEY 2
A. Fertile vesicle producing many sporangiola covered with capitate appendages; homothallic, zygosporangium smooth and thin; suspensors with appendages—— Radiomyces
AA. Fertile vesicle producing a single sporangiole covered with acicular appendages; presumably heterothallic; zygospores unknown—— Hesseltinella
KEY 3
A. Sporophores verticellately branched—— Chaetocladium
AA. Sporophores dichotomously branched—— Dichotomocladium
KEY 4
A. Terminal sporangia always accompanied by laterally produced sporangiola——B
AA. Either uni- or multispored sporangiola produced; terminal sporangia present or not——G
B. Sporangiola vary from uni- to multispored on the same sporophore—— Backusella
BB. Sporangiola always multispored——C
C. With a more or less conspicuous subsporangial swelling—— Fennellomyces
CC. Without a subsporangial swelling——D
D. Sporangium and/or sporangiole apophystate—— Thamnostylum
DD. Sporangium nonapophysate, sporophore constricted beneath the sporangium——E
E. Sporangiola formed laterally on twisted, contorted sporophoric branches; sporangiolar columella cylindrical to hour-glass shaped—— Pirella
EE. Sporangiola formed on fertile branches that are more dichotomous or more or less verticellately arranged on the sporophore; sporangiolar columellae globose to obovoid or compressed——F
F. Fertile branches several times dichotomously branched; spines not formed—— Thamnidium
FF. Sporangiolar pedicels simple, more or less verticellately or irregularly arranged on fertile branch; sterile spines formed—— Helicostylum
G. Pedicellate sporangiola produced on a conspicuous vesicle, uni- or multispored——H
GG. Pedicellate sporangiola arise directly from sporophore or its branches, or inconspicuous vesicle present at point where several vesicles arise——K
H. Sporangiola unispored—— Phascolomyces
HH. Sporangiola multispored——I
I. Vesicles intercalary—— Thamnostylum
II. Vesicles terminal——J
J. Vesicles globose to obovoid; sporophores unbranched; sporangiola globose—— Cokeromyces
JJ. Vesicles irregular; sporophores usually branched; sporangiola obpyriform—— Zychaea
K. Sporangiola globose to subglobose, nonapophysate or pedicel constricted beneath sporangiole——L
KK. Sporangiola obovoid to pyriform, apophysate——N
L. Branching verticellate; sterile spines present or not——O
LL. Branching sympodial, bi- or trifurcate, subdichotomous or dichotomous; sterile spines lacking——M
M. Sporangiola borne on dichotomous branches——_Thamnidium_
MM. Sporangiola borne on sympodial to bi- or trifurcate, or subdichotomous branches—— Ellisomyces
N. Columellae of sporangiola usually large, obpyriform, cylindrical to hour-glass shaped—— Pirella
NN. Columellae of sporangiola small, hemisphaerical—— Thamnostylum repens
O. Sterile spines formed; pedicel more or less straight or apically curved, may or may not arise from an small enlargement, arise verticellately or irregularly; best growth below 20 C—— Helicostylum
OO. Sterile spines lacking; pedicels recurved to twisted and contorted, arise verticellately, often from a small vesicle; best growth below 20 C—— Kirkomyces
KEY 5
A. Fertile vesicle globose to obovoid; pedicels long, curved to twisted and contorted; denticles and sporangiola monomorphic—— Benjaminiella
AA. Fertile vesicle cylindrical; pedicels short and straight; denticles and sporangiola dimorphic—— Mycotypha
KEY 6
A. Thermophilic—— Thermomucor
AA. Mesophilic——B
B. Sporangium dumbbell shaped—— Halteromyces
BB. Sporangium globose to ovoid——C
C. Apophysis constricted below the sporangium which is globose—— Gongronella
CC. Apophysis not obviously constricted, but if slightly constricted then sporangium obovoid——D
D. Large, terminal, dark, clavate to fusiform chlamydospores developed by aerial hyphae—— Chlamydoabsidia
DD. Chlamydospores, if present, hyaline, inconspicuous——E
E. Stolons absent——F
EE. Stolons present; sporangial branches more or less straight or if circinate developed opposite rhizoids——G
F. Sporangia bearing branches more or less straight; sporangial wall deliquescent; homothallic; zygosporangial wall ornamented—— Protomycocladus
FF. Sporangial bearing branches circinate; sporangial wall persistent; heterothallic or presumably so; zygosporangial wall more or less smooth—— Circinella
G. Sporangial branches in umbels near apex of sporophore——_Rhizopodopsis_
GG. sporangial branches verticellate or if umbellate developed opposite rhizoids——H
H. Some sporangia formed opposite rhizoids——I
HH. Sporangia formed on aerial hyphae or on stolons, never opposite rhizoids——J
I. Sporangial wall deliquescent; some apophyses slightly swollen; columella not collapsing; sporangiospores smooth walled—— Apophysomyces
II. Sporangial wall dry, evanescent (fractures), appears persistent; apophysis not swollen; columella collapsing; sporangiospore wall spinose or striate—— Rhizopus
J. Growth good at 37 C; aerial mycelium determinant; sporophores borne randomly on stolons; optimal temperature for zygospore formation 31 C; suspensors without appendages—— Mycocladus
JJ. Growth poor or lacking at 37 C; aerial mycelium indeterminant; sporophores usually borne in umbels or verticels on stolons; optimum temperature for zygospore formation 26 C; one or both suspensors with appendages—— Absidia
KEY 7
A. Sporophores unbranched—— Spinellus
AA. Sporophores branched——B
B. Sporophores umbellately branched initially, final branch dichotomous; sterile spinelike branches produced; parasitic on insects in the tropics—— Sporodiniella
BB. Sporophores sympodial to dichotomous; sterile spinelike branches not formed; parasitic on mushrooms, generally in temperate regions——C
C. Sporophores dichotomously branched; all sporangia more or less the same size; grows well at 25 C; suspensors slightly unequal—— Syzygites
CC. Sporophores simple, sympodial or dichotomous; sporangia of various sizes; grows well at 19 C and below; suspensors strongly unequal—— Dicranophora
KEY 8
A. Sporophore constricted below the sporangium, without subsporangial swelling or trophocyst—— Pilaira
AA. Sporophore with a well-developed subsporangial vesicle; trophocysts produced——B
B. Subsporangial vesicle arises a short distance below the sporangium, which is not forcibly released; grows well on ordinary laboratory media—— Utharomyces
BB. Subsporangial swelling arises immediately below the sporangium, which is forcibly released; usually requires addition of ferrichrome, hemin, or dung extract for growth on laboratory culture media—— Pilobolus
KEY 9
A. Thermophilic—— Rhizomucor (in part, R. meihei, R. pusillus, R. tauricus )
AA. Mesophilic——B
B. Rhizoids and stolons usually formed——C
BB. Rhizoids and stolons not formed, sporangia not borne verticellately——D
C. Smaller sporangia borne verticellately below a larger terminal sporangium—— Actinomucor
CC. All sporangia more less the same, not subtended by smaller sporangia—— Rhizomucor (in part, R. endophyticus, R. variabilis )
D. In nature a facultative, gall-forming parasite of other Mucorales; suspensors with short projections—— Parasitella
DD. In nature saprobic; suspensors without projections——E
E. Sporophores arising from substrate mycelium——F
EE. Sporophores arising from aerial hyphae—— Hyphomucor
F. Sporophores highly branched with many, usually small sporangia (usually less than 80 μm in diam); zygosporangial wall ornamentated with stellate projections, reddish-brown in color—— Circinomucor
FF. Sporophores simple or occasionally branched, some sporangia over 80 μm in diam; zygosporangial ornaments not stellate, wall brown to black——G
G. Sporangia with a clavate to obclavate, relatively large columella; sporangiospores usually ellipsoid; hetero- or homothallic; zygosporangial wall ornamentation verrucose; suspensors subunequal—— Mucor
GG. Sporangia with relatively small, sphaerical or oblate columellae; sporangiospores ellipsoidal, sphaerical, or bacilliform; always homothallic; zygosporangial wall with spinose ornaments; suspensors distinctly unequal—— Zygorhynchus
Synopsis of Genera
Note: More information about each genus can obtained by consulting the latest version of Ainsworth & Bisby’s Dictionary of the Fungi (Kirk et el., 2001).
ABSIDIA van Tieghem, 1878, subgen Absidia (Ann. Sci. Nat. Bot., Sér. 6, 4: 350); 26+ spp. (Ellis and Hesseltine, 1965 —ILLUS. and KEY TO SPP., 1966 —ILLUS.; Hesseltine and Ellis, 1961 —illus.; 1964 —ILLUS. and KEY TO SPP., 1966 —ILLUS. and KEY TO SPP.; Vanova, 1968 —illus. and key to spp., 1969, 1971 —illus., 1983 —ILLUS. and KEY TO SPP.; Milko, 1970b —illus. and key to spp.; Schipper, 1990 —key; Hesseltine et al., 1990 —key; Chen and Zheng, 1998 —thermophile).
= Tieghemiella Berl. & de Toni, 1888 (In Saccardo, Syll. Fung. 7: 215).
= Lichthemia Vuillemin, 1903 (Bull. Soc. Mycol. France 19: 126).
= Proabsidia Vuillemin, 1903 (Bull. Soc. Mycol. France 19: 116).
= Pseudoabsidia Bainier, 1903 (Bull. Soc. Mycol. France 19: 155).
= Pseudoabsidia Saccardo, 1905 (Syll. Fung. 17: 504; fide Kirk, 1985)
= Protoabsidia Naumov, 1935 [Opredelitel’ Mukorovykh (Mucorales), Ed. 2, pp. 23 and 78; nomen nudum, without a Latin diagnosis, Art. 36.1 of the ICBN, Greuter et al., 2000].
ACTINOMUCOR Schostakowitsch, 1898 (Ber. Deutsch. Bot. Ges. 16: 155); 2 spp. (C. R. Benjamin and Hesseltine, 1957 —illus.; Zycha et al., 1969 —ILLUS.; Jong and Yuan, 1985 —illus.).
= Glomerula Bainier, 1903 (Bull. Soc. Mycol. France 19: 154).
[ Actinomotierella, see Mortierellaceae (Mortierellales)]
APOPHYSOMYCES Misra, 1979 (in Misra, Srivasatava and Lata, Mycotaxon 8: 377); 1 sp. (Misra et al. 1979 —illus.)(a synonym of Absidia according to O’Donnell, 1979; a human pathogen according to Ellis and Ajello 1982 [also report a method to induce sporulation in Apophysomyces ]; Padhyay and Ajello, 1988-sporulation).
BACKUSELLA Hesseltine & Ellis, 1969 (In Ellis and Hesseltine, Mycologia 61: 863); 3 spp. (Ellis and Hesseltine, 1969 —illus.; Benny and Benjamin, 1975 —ILLUS. and KEY TO SPP.; Stalpers and Schipper, 1980 —illus.).
BENJAMINIELLA von Arx, 1981 (_Genera of Fungi Sporulating in Pure Culture_, 3rd Ed., p. 60); 2 spp. (Benjamin, 1960 —ILLUS. as Cokeromyces poitrasii R. K. Benjamin; Benny and Benjamin, 1976 —ILLUS. as Mycotypha poitrasii (Benjamin) Benny & Benjamin; Brain and Young, 1979 —illus. as M. poitrasii; Cole and Samson, 1979 —illus. as M. poitrasii; Benny et al., 1985 —ILLUS. and key to spp.; Kirk, 1989 —ILLUS. and KEY TO SPP.).
= Benjaminia Pidoplichko & Mil’ko, 1971 ( Atlas of Mucoralean Fungi, p. 96) (see Benny and Benjamin, 1976).
BLAKESLEA Thaxter, 1914 [Bot. Gaz. (Crawfordsville) 58: 353]; 2 spp. (Thaxter, 1914 —ILLUS.; B. S. Mehrotra and Baijal, 1968 —illus.; Benny and O’Donnell, 1978a —illus.; Kirk, 1984 —ILLUS. and KEY TO SPP.; Zheng and Chen, 1986).
CHLAMYDOABSIDIA Hesseltine & Ellis, 1966 (Mycologia 58: 761); 2 spp (Hesseltine and Ellis, 1966—illus.; Behera and Mukerji, 1974 —illus.)[transferred to Absidia by Milko, 1970a].
CHAETOCLADIUM Fresenius, 1863 (Beitr. Mykol., p. 97); 2 spp. (Benny and Benjamin, 1976 —ILLUS. and KEY TO SPP.).
CHOANEPHORA Currey, 1873 (J. Linn. Soc., Bot. 13: 578); 2 spp. (Thaxter, 1903 —ILLUS.; Poitras, 1955 —illus.; Hesseltine and Benjamin, 1957 —illus.; Kirk, 1984 —ILLUS. and KEY TO SPP.; Sakai et al., 2000-pollination).
= Cunninghamia Currey, 1873 [J. Linn. Soc., Bot. 13: 334; non Cunninghamia R. Br., 1826 (In Richard, Comm. Bot. Conif. Cycad, p. 149)].
= Choanephorella Vuill., 1904 (Bull. Soc. Mycol. France 20: 28; nomen nudum, Art. 41.1 of the ICBN (Greuter et al., 2000).
CIRCINELLA van Tieghem & Le Monn., 1873 (Ann. Sci. Nat. Bot., Sér. 5, 17: 298); 9 or 10 spp. (Hesseltine and Fennell, 1955 —ILLUS. and KEY TO SPP.; Hesseltine and Ellis, 1961 —illus.; Faurel and Schotter, 1965 —illus.; Milko, 1969 —added species of Pirella; Patil and Kale, 1981 —illus.; Naganishi, 1974; Arambarri and Cabello, 1996).
= Circinumbella van Tieghem & Le Monn., 1872 (Compt. Rend. Hebd. Séances Acad. Sci. 74: 999).
CIRCINOMUCOR v. Arx, 1983 (Sydowia 35: 17); 9 spp. (Schipper, 1976 —illus. as Mucor spp.; Von Arx, 1982 —key to spp. —a synonym of Mucor).
[ Coemansia, see Kickxellaceae (Kickxellales)]
COKEROMYCES Shanor, 1950 (In Shanor, Poitras, and Benjamin, Mycologia 42: 272); 1 sp. (Benny and Benjamin, 1976 —ILLUS.; Jeffries and Young, 1983a —illus., 1983b —illus.).
CUNNINGHAMELLA Matruchot, 1903 (Ann. Mycol. 1: 47); 5 spp. [10 spp. + 2 varieties to 3 spp. + 1 var. recognized] ( Baijal and Mehrotra, 1980 —ILLUS. and KEY TO SPP; Weitzman, 1984; Liu et al., 2001-phylogeny; Zheng and Chen, 2001 —monograph).
= Actinocephalum Saito, 1905 [Bot. Mag. (Tokyo) 19: 36].
= Saitomyces Ricker, 1906 (J. Mycology 12: 61).
= Muratella Bainier & Sartory, 1913 (Bull. Soc. Mycol. France 29: 129).
DICHOTOMOCLADIUM Benny & R. K. Benjamin, 1975 (Aliso 8:338); 5 spp. (Benny and Benjamin, 1975 —ILLUS. and KEY TO SPP.; 1993—ILLUS. and KEY TO SPP.).
DICRANOPHORA Schröter, 1886 (Ber. Schles. Ges. vaterl. Kult. 64:184); 1 sp. (Dobbs, 1938 —ILLUS.; Voglmeyr and Krisai-Greilhuber, 1996 —ILLUS.).
[ Dimargaris, see Dimagaritaceae (Dimargaritales)]
[ Dipsacomyces, see Kickxellaceae (Kickxellales)]
[ Dispira, see Dimagaritaceae (Dimargaritales)]
[ Dissophora, see Mortierellaceae (Mortierellales)]
[ Echinosporangium, see Mortierellaceae (Mucorales)].
ELLISOMYCES Benny & R. K. Benjamin, 1975 (Aliso 8: 330); 1 sp. (Benny and Benjamin, 1975 —ILLUS.).
FENNELOMYCES Benny & R. K. Benjamin, 1975 (Aliso 8: 328); 4 spp. (Benny and Benjamin, 1975 —ILLUS.; Misra et al., 1979 —illus.; Mirza et al., 1979 —illus.).
[ Endogone, see Endogonaceae (Endogonales])
GILBERTELLA Hesseltine, 1960 (Bull. Torrey Bot. Club 87: 24); 1 sp. (Hesseltine, 1960a —illus.; O’Donnell et al., 1977a —illus., 1977c —illus.; Benny, 1991 —ILLUS.).
GONGRONELLA Ribaldi, 1952 (Riv. Gen. Biol., N.S., 44:164); 2 spp. (Hesseltine and Ellis, 1961, 1964).
HALTEROMYCES Shipton & Schipper, 1975 (Antonie van Leeuwenhoek Ned. Tijdschr. Hyg. 41: 337); 1 sp. (Shipton and Schipper, 1975 —illus.).
[ Helicocephalum, see Helicocephalidaceae (Zoopagales)]
HELICOSTYLUM Corda, 1842 (_Icon. Fung_. 5: 18); 2 spp. (Upadhyay, 1973 —illus. and key to spp.; Benny, 1995b —ILLUS. and KEY TO SPP.).
= Haynaldia Schulzer von Müggenberg, 1866 (Verh. Zool.-Bot. Ges. Wien 17: 37; fide Upadhyay, 1973).
= Bulbothamnidium Klein, 1870 (Verh. Zool.-Bot. Ges. Wien 20: 557; fide Upadhyay, 1973).
= Chaetostylum van Tiegh. & Le Monn., 1873 (Ann. Sci. Nat. Bot., Sér. 5, 17: 328; fide Upadhyay, 1973)
HESSELTINELLA Upadhyay, 1970 (Persoonia 6: 111); 1 sp. (Benny and Khan, 1988; Benny and Samson, 1989; Benny and Benjamin, 1991 —monograph).
HYPHOMUCOR Schipper & Lunn, 1986 (In Schipper, Mycotaxon 27: 83); 1 sp. (Schipper, 1986b —illus.).
[ Kickxella, see Kickxellaceae (Kickxellales)]
KIRKOMYCES Benny, 1995 (Mycologia 87:922); 1 sp. (The correct name for Kirkia Benny, a later homonym, Benny, 1995a) (Benny, 1995b —ILLUS.).)
= Kirkia Benny, 1995 (Mycologia 87:261); 1 sp. (Benny, 1995b —ILLUS.).
[ Linderina, see Kickxellaceae (Kickxellales)]
[ Martensella, see Kickxellaceae (Kickxellales)]
[ Martensiomyces, see Kickxellaceae (Kickxellales)]
[ Micromucor, a synonym of Umbelopsis (Umbelopsidaceae, Mucorales)]
[ Modicella, see Mortierellaceae (Mortierellales)]
Mortierella, see Mortierellaceae (Mortierellales)]
MUCOR Micheli ex L.: Fr., 1832 [Syst. Mycol. 3: 317; nomen conserv., Art. 13.1(d) of the ICBN (Greuter et al., 2000; see Korf, 1982, 1983)]; 20 to 42 + spp. (Schipper, 1973 —ILLUS., 1975 —ILLUS., 1976 —ILLUS., 1978a —ILLUS. and KEY TO SPP. of Mucor s.l.; Schipper et al., 1975 —illus.; Michaelides and Spotts, 1979 —review Mucor piriformis disease of pear; Stalpers and Schipper, 1980 —illus.; B. S. Mehrotra and B. M. Mehrotra, 1978 —illus.; Subrahmanyam, 1983 —illus.; Kirk, 1986 —proposal to conserve Mucor Fres. over Mucor Micheli ex L.: Fr.; Chen and Zheng, 1986; Schipper, 1989; Schipper and Samson, 1994; Watanabe, 1994; Zalar et al., 1997).
= Mucor Micheli, 1729 ( Nova Plantarum Genera, p. 215; fide Kirk, 1986).
= Mucor Micheli ex Linnaeus, 1753 ( Species Plantarum 2: 1185; fide Kirk, 1986).
= Hydrophora Tode: Fr., 1832 ( Syst. Mycol. 3: 313; fide Kirk, 1986); nomen conserv., Art. 13.1(d) of the ICBN (Greuter et al., 2000; see Korf, 1982, 1983)].
= Hydrophora Tode, 1791, p.p. ( Fungi Mecklenbergensis selecti 2: 5; fide Hesseltine, 1955).
= Chionyphe Thienemann, 1839 (Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 19: 19; fide Hesseltine, 1955).
= Calyptromyces H. Karsten, 1849 [Bot. Zeitung (Berlin) 7: 365; fide Hesseltine, 1955].
= Mucor Fresenius, 1850 ( Beiträge zur Mykologie, p. 7; fide Kirk, 1986).
= Pleurocystis Bonorden, 1851 ( Handb. Allge. Mykol., p. 124; fide Hesseltine, 1955).
= Scitovszkya Schulzer von Müggenberg, 1866 (Verh. Zool. -Bot. Ges. Wien 16: 36; fide Hesseltine, 1955).
= Chlamydomucor Brefeld, 1889 (Unters. Ges. Mycol. 8: 223)(Ellis et al., 1974, 1976).
?= Ascidiophora Reich., 1901 (fide Saccardo, Syll. Fung. 15: 48).
MYCOCLADUS Beauverie, 1900 (Ann. Univ. Lyon, Sér. 2, Sciences, Médicine 3: 162); 6 spp. (Mirza et al., 1979 —key to spp.)(considered a subgenus of Absidia by Hesseltine and Ellis, 1964).
= Absidia van Tiegh. subgen. Mycocladus (Beauv.) Hesseltine & Ellis, 1964 (Mycologia 56: 569).
MYCOTYPHA Fenner, 1932 (Mycologia 24: 196); 3 spp. (Benny and Benjamin, 1976 —ILLUS and key to spp.; Brain and Young, 1979 —illus.; Benny et al., 1985 —ILLUS. and KEY TO SPP.; Edelmann and Klomparens, 1995 —ultrastructure).
PARASITELLA Bainier, 1903 (Bull. Soc. Mycol. France 19: 153); 1 spp. (Schipper, 1978b —illus.).
PHASCOLOMYCES Boedijn, l959 (Sydowia 12: 349); 1 sp. (Benny and Benjamin, 1976 —ILLUS.; Jeffries and Young, 1978 —illus.).
PHYCOMYCES Kunze: Fr., 1821 [Syst. Mycol. 3: 308; nomen conserv., Art. 13.1(d) of the ICBN (Greuter et al., 2000; see Korf, 1982, 1983)]; 3 spp. (C. R. Benjamin and Hesseltine, 1959—illus. and KEY TO SPP.); O’Donnell et al., 1976 —illus., 1978b —illus.; Cerdá-Olmedo and Lipson, 1987 —Review).
= Phycomyces Kunze, 1823 (In Kunze and Schmidt, Mykol. Hefte 2: 113).
PILAIRA van Tieghem, 1875 (Ann. Sci. Nat. Bot., Sér. 6, 1: 51); 5 spp. (Grove, 1934 —ILLUS. and KEY TO SPP.; Mil’ko, 1970b —illus.).
PILOBOLUS Tode: Fr., 1832 (Syst. Mycol. 3: 312); 10 to 13 spp. + 2 varieties (Grove, 1934 —ILLUS. and KEY TO SPP.; McVickar, 1942 —illus.; Lyr, 1954 —illus.; Hu et al., 1989 —revision)
= Pilobolus Tode, 1784 (In Schrift. Berl. Gesell. naturf. Fr., 5: 46).
= Pycnopodium Corda, 1842 (_Icon. Fung_. 5: 18; fide Hesseltine, 1955).
[ Piptocephalis, see Piptocephalidaceae (Zoopagales)]
PIRELLA Bainier, 1882 ( Étude sur les Mucorinées, p. 83); 2 spp. (Hesseltine, 1960b —illus.; Benny and Schipper, 1992 —ILLUS. and KEY TO SPP.).
= Mucor Micheli ex L.: Fr. subgen. Pirella (Bain.) Schröt., 1893 [In Engler & Prantl, Die naürl. Pflanzenfam. 1(1): 125; fide Hesseltine, 1960b].
= Mucor Micheli ex L.: Fr. sect. Piromyces Naumov, 1935
[ Opredel. Mukoroykh ( Mucorales ), Ed. 2, p. 29; nomen nudum, without a Latin diagnosis, Art. 36.1 of the ICBN (Greuter et al., 2000); fide Hesseltine, 1960b].
POITRASIA Kirk, 1984 (Mycol. Papers 152: 51); 1 sp. (Hesseltine and Benjamin, 1957 —illus. as Choanephora circinans; Kirk, 1984 —ILLUS.).
PROTOMYCOCLADUS Schipper & Samson, 1994 (Mycotaxon 50:487); 1 sp. (Schipper and Samson, 1994—illus.).
RADIOMYCES Embree, 1959 (Amer. J. Bot. 46: 25); 3 spp. (Benny and Khan, 1988; Benny and Samson, 1989 —illus.; Benny and Benjamin, 1991 —revision).
= Radiomycopsis Pidopl. & Mil’ko, 1971 (Atlas of Mucoralean Fungi, p. 95).
RHIZOMUCOR (Lucet & Cost.) Wehmer ex Vuill., 1931 (_Les Champignons parasites et les mycoses de l’hommes_, p.19); 5 spp. (Schipper, 1978b —illus.; Zheng and Chen, 1991, 1993 —key; Zheng and Jiang, 1995 —endophyte).
= Mucor Micheli ex L.: Fr. sect. Rhizomucor Lucet & Cost., 1899 (Compt. Rend. Hebd. Séances Acad. Sci 129: 1033; fide Schipper, 1978b).
= Mucor Micheli ex L.: Fr. subgen. Rhizomucor (Lucet & Cost.) Sacc. & Syd. 1902 (In Saccardo, Syll. Fung. 16: 385; fide Schipper, 1978b).
= Rhizomucor (Lucet & Cost.) Wehmer, 1907 (In Lafar, Handb. Techn. Mykol. 4: 459; fide Schipper, 1978b).
RHIZOPODOPSIS Boedijn, 1959 (Sydowia 12: 330); 1 sp. (Boedijn, 1959 —illus. [also illus. by O’Donnell, 1979]).
RHIZOPUS Ehrenberg, 1820 (Nova Acta Phys.-Med. Acad. Caes. – Leop. – Carol. Nat. Cur. 10: 198); 4 to 10 spp. (Schipper, 1984 —ILLUS. and KEY TO TAXA; Schipper and Stalpers, 1984 —ILLUS. and KEY TO TAXA; Yuan and Jong, 1984 —illus.; Yuan et al., 1985 —illus.; Kirk, 1986 —proposal to conserve Rhizopus; Schipper and Samson, 1994; Weitzman et al., 1996; Zheng and Chen, 1998).
= Ascophora Tode: Fr., 1832 [Syst. Mycol. 3(2): 309; fide Kirk, 1986), nomen concerv., Art. 13.1(d) of the ICBN; Greuter et al., 2000 (see Korf, 1982, 1983). (_Ascophora_ Tode, 1790 (_Fungi Mecklenbergensis selecti_ 1: 13; fide Kirk, 1986)
= Amylomyces Calmette, 1892 (Ann. Inst. Pasteur 6: 611; fide Elliset al., 1974); see also Ellis et al. (1976).
[ Rhopalomyces, see Helicocephalidaceae (Zoopagales)]
SAKSENAEA Saksena, 1953 (Mycologia 45: 434); 1 sp. (Saksena, 1953 —ILLUS.; Ajello et al., 1976 — illus. and first report as a human pathogen; Ellis and Ajello, 1982 — technique for inducing sporulation in Saksenaea; Padhyay and Ajello, 1988 — technique for inducing sporulation in Saksenaea ).
[ Sigmoideomyces, see Cunninghamellaceae (Mucorales)].
[ Spinalia, see Dimagaritaceae (Dimargaritales)]
SPINELLUS van Tieghem, 1875 (Ann. Sci. Nat., Bot., S�r. 6, 1:66); 5 spp. (Indoh, 1961 —illus.; Ellis and Hesseltine, 1962; Watson, 1964 —illus.; Zycha et al., 1969 —illus. and KEY TO SPP.; Strid, 1974; O’Donnell, 1979 —illus.; Overton, 1997)
[ Spirodactylon, see Kickxellaceae (Kickxellales)]
[ Spiromyces, see Kickxellaceae (Kickxellales)]
[ Sporodinia, a synonym of Syzygites (Mucoraceae, Mucorales)].
SPORODINIELLA Boedijn, 1958 (Ann. Mycol. 12:336); 1 sp. (Evans and Samson, 1977 —ILLUS.; Gbaja and Young, 1985 —illus.; Chien and Hwang, 1997).
SYNCEPHALASTRUM Schröter, 1886 (In Cohn’s Kryptogamen-Flora von Schlesen 3(2): 217); 2 sp. (Benjamin, 1959 —ILLUS.; Zycha et al., 1969 —illus.; Misra, 1975 —illus.; Benjamin and Tucker, 1978 —ILLUS.; Schipper and Stalpers, 1983 —illus.; Zheng et al., 1988 —ILLUS.).
[Syncephalis, see Piptocephalidaceae (Zoopagales)]
SYZYGITES Ehrenberg, 1818 ( Sylvae Mycol. Berol., p. 25); 1 sp. (Hesseltine, 1957; Benny and O’Donnell, 1978b —illus.; O’Donnell, 1979 —illus.; Ekpo and Young, 1979 —illus.; Kovaks and Sundberg, 1999).
THAMNIDIUM Link, 1809 (Ges. Naturk. Freunde Berlin Mag. Neuesten Entdeck. Gesammten 3: 31); 1 sp. (Benny, 1973 —ILLUS.).
= Thamnidium Link ex Gray, 1821 ( A Natural Arrangement of British Plants 1: 560; fide Benny, 1973).
= Thamnidium Link ex Wallroth, 1833 [ Flora Crypt. Germ. 4(2): 324; fide Benny, 1973].
= Melidium Eshweiler: Fr., 1832 (Syst. Mycol. 3: 330; fide Benny, 1973), nomen conserv., Art. 13.1(d) of the ICBN (Greuter et al., 2000; see Korf, 1982, 1983).
= Melidium Eshweiler, 1822 ( De Fruct. Gen. Rhizomorphae Comment., p. 33; fide Benny, 1973).
[ Thamnocephalis, see Cunninghamellaceae (Mucorales)].
THAMNOSTYLUM v. Arx & Upadhyay, 1970 (In von Arx, Genera of Fungi Sporulating in Pure Culture, p. 247); 4 spp. (Upadhyay, 1973 —illus. and key to spp.; Benny and Benjamin, 1975 —ILLUS. and KEY TO SPP.).
THERMOMUCOR Subrahamanyam, Mehrotra & Thirumalacher, 1977 (Georgia J. Sci. 35: 1); 1 sp. (Subrahamanyam et al., 1977 —illus.; Schipper, 1979 —ILLUS.).
[ Tieghemiomyces, see Dimagaritaceae (Dimargaritales)]
[ Umbelopsis, see Umbelopsidaceae (Mucorales)]
UTHAROMYCES Boedijn, 1959 (Sydowia 12: 340); 1 sp. (Kirk and Benny, 1980 —ILLUS.).
ZYCHAEA Benny & R. K. Benjamin, 1975 (Aliso 8: 334); 1 sp. (Benny and Benjamin, 1975 —ILLUS.).
ZYGORHYNCHUS Vuillemin, 1903 (Bull. Soc. Mycol. France 19: 116); 7 spp. (Hesseltine et al., 1959 —illus. and key to spp.; Schipper et al., 1975 —illus.; O’Donnell et al., 1978 —illus.; Schipper, 1986a —illus. and KEY TO SPP., Zheng, 2002 —key to species).
Updated Feb 10, 2005