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 that 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 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. Thamnidiaceae is considered a synonym of Mucoraceae because the type for the latter family, Thamnidium elegans Link, is now a member of the Mucoraceae as presently circumscribed.
A data set consisting of six taxa from the Mortierellales and 75 spp. containing at least one species all of the genera in culture of the Mucorales (tef-1a, 18S rDNA, 28S rDNA, morphology) of O’Donnell et al. (2001) showed that the Mucoraceae clade had 91% bootstrap support (BP). Voigt and Olsson (2008) analyzed a data set (act, tef-1a, 18S rRNA, 28S rRNA) containing selected species of 50 genera of the Mucorales by neighbor-joining (100% BP) and strict consensus (98% BP) for the Mucoraceae
Absidiaceae was considered a synonym of Mucoraceae and Dicranophoraceae was treated as a synonym of Thamnidiaceae by Hawksworth et al. (1995) and Kirk et al. (2001). All of the remaining families of the Mucorales were being treated as synonyms of Mucoraceae based on the phylogenetic study of the Mucorales published by O’Donnell et al. (2001) in a previous version (10 February 2005 to 30 July 2009) of this website (http://www.zygomycetes.org).
Thamnidiaceae is considered a synonym of Mucoraceae because the type for the latter family, Thamnidium elegans Link, is now a member of the Mucoraceae as presently circumscribed. Mucoraceae contains five genera here based on the above phylogenetic analyses (Helicostylum, Mucor s.s., Pilaira, Pirella, Thamnidium). The other genera that constituted the Thamnidiaceae sensu Benny (Benny, 1973; Benny and Benjamin, 1975, 1976) included Backusella Hesselt. & J.J. Ellis, Chaetocladium Fres., Cokeromyces Shanor, Dichotomocladium Benny & R.K. Benj, Ellisomyces Benny & R.K. Benj, Fennellomyces Benny & R.K. Benj, Mycotypha Fenner, Phascolomyces Boedijn, Thamnostylum v. Arx & Upadhyay, and Zychaea Benny & R.K. Benj. These fungi can be found in other families (Chaetocladiaceae — Chaetocladium, Mycotyphaceae — Mycotypha) of the Mucorales or the Synopsis of Genera with Unknown Affinities (Backusella, Dichotomocladium, Elllisomyces, Fennellomyces, Phascolomyces, Thamnostylum, Zychaea).
Classification
Mucoraceae Dumortier 1822 (Com. Bot., p. 69 et 82).
= Thamnidiaceae Fitzpatrick, 1930 (The Lower Fungi Phycomycetes, p. 253).
Saprobes. Mycelium well developed; rhizoids and stolons lacking. Sporangiophores erect or ascending, simple or branched, arising from substrate mycelium or aerial hyphae. Sporangia similar, more or less globose, columellate, multispored, without an apophysis; wall diffluent, deliquescent, or persistent. Sporangiola, if formed, similar, more or less globose and nonapophysate or obpirform and apophysate, columellate, wall persistent. Suspensors formed on aerial hyphae or sporangiophores or aerial hyphae, opposed orapposed, more or less equal, 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 or spines, , 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 or apposed, more or less equal, unadorned.
Type genus: Mucor s.s.
KEY TO THE GENERA OF MUCORACEAE
A. Sporangia and sporangiola produced —— B.
AA. Sporangia produced —— D
B. Sporangiola obpiriform, 1-3 or 4 produced on a relatively long, twisted and contorted lateral branch arising from the sporangiophore —— Pirella
BB. Sporangiola more or less globose, many produced on fertile branches that are straight, curved or dichotomously branched —— C
C. Fertile branches terminate in sterile spines, arise pseudovertilcellately from the main sporangiophore or its branches —— Helicostylum
CC. Fertile branches do not terminate in sterile spines, arise verticellately from the main sporangiophore or its branches, remainder of branching dichotomous —— Thamnidium
D. Sporangiophores simple or sympodially branched; sporangia more or less globose, off-white to grayish in color; sporangiolar wall deiquescent —— Mucor s.s.
DD. Sporangiophores usually unbranched; sporangia broadly ellipsoid; usually dark when mature; sporangiolar wall perisitent —— Pilaira
Synopsis of Genera in the Mucoraceae
Note: More information about each genus can obtained by consulting the latest version of Ainsworth & Bisby’s Dictionary of the Fungi (Kirk et el., 2008), and under each genus in this website (http://www.zygomycetes.org)
HELICOSTYLUM Corda, 1842 (Icon. Fung. 5: 18); 2 spp. (Upadhyay, 1973 — illus. and key to spp.; Benny, 1995 — 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)
MUCOR Fresenius, 1850 [Beiträge zur Mykologie, p. 7; nomen conserv., Art. 14.1of the ICBN (McNeill et al., 2006; fide Kirk, 1986); 20 to 42 + spp. (Schipper, 1973 — ILLUS., 1975 — ILLUS., 1976 — ILLUS., 1978 — 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.; Mehrotra and 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 (McNeill et al., 2006; 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 Micheli, 1729 (Nova Plantarum Genera, p. 215; fide Kirk, 1986).
= Mucor Micheli ex L.: Fr., 1832 [Syst. Mycol. 3: 317; nomen sanct., Art. 13.1(d) of the ICBN (McNeill et al., 2006; see Korf, 1982, 1983)];
= 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).
?= Circinomucor von Arx, 1982 [1984] (Sydowia 35:17) fide (http://www.indexfungorum.org/Names/Names.asp/; 18 November 2009).
PILAIRA van Tieghem, 1875 (Ann. Sci. Nat. Bot., Sér. 6, 1: 51); 5 spp. (Grove, 1934 — ILLUS. and KEY TO SPP.; Mil’ko, 1970-illus.; Zheng and Liu, 2009 — ILLUS., new spp.).
PIRELLA Bainier, 1882 (Étude sur les Mucorinées, p. 83); 2 spp. (Hesseltine, 1960 — illus.; Benny and Schipper, 1992 — ILLUS. and KEY TO SPP.).
= Mucor Micheli ex L.: Fr. subgen. Pirella (Bain.) Schröt., 1893 [In Engler & Prantl, Die natürl. Pflanzenfam. 1(1): 125; fide Hesseltine, 1960].
= 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 (McNeill et al., 2006); fide Hesseltine, 1960].
THAMNIDIUM Link, 1809 (Ges. Naturk. Freunde Berlin Mag. Neuesten Entdeck. Gesammten 3: 31); 1 sp. (Benny, 1973 — ILLUS., 1992 — ILLUS..).
= Thamnidium Link ex Gray, 1821 (A Natural Arrangement of British Plants 1: 560; fide Benny, 1973, 1992).
= Thamnidium Link ex Wallroth, 1833 [Flora Crypt. Germ. 4(2): 324; fide Benny, 1973, 1992].
= __Melidium_ Eshweiler: Fr., 1832 (Syst. Mycol. 3: 330; fide Benny, 1973, 1992), nomen conserv., Art. 13.1(d) of the ICBN (McNeill et al., 2006; see Korf, 1982, 1983).
= Melidium Eshweiler, 1822 (De Fruct. Gen. Rhizomorphae Comment., p. 33; fide Benny, 1973, 1992).
Genera Not Accepted Here but Included in Mucoraceae by Some Authors
[Absidia — see Absidiaceae (Mucorales)].
[Actinomucor — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Actinomotierella — see Mortierellaceae (Mortierellales)]
[Apophysomyces — see Saksenaeaceae (Mucorales)].
[Backusella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Benjaminiella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Blakeslea — see Choanephoraceae (Mucorales)].
[Chlamydoabsidia — see Absidiaceae (Mucorales)].
[Chaetocladium — see Chaetocladiaceae (Mucorales)].
[Choanephora — see Choanephoraceae (Mucorales)].
[Circinella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Circinomucor — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Coemansia — see Kickxellaceae (Kickxellales)]
[Cokeromyces — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Cunninghamella — see Cunninghamellaaceae (Mucorales)].
[Dichotomocladium — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Dicranophora — see Dicranophoraceae (Mucorales)].
[Dimargaris — see Dimargaritaceae (Dimargaritales)]
[Dipsacomyces — see Kickxellaceae (Kickxellales)]
[Dispira — see Dimagaritaceae (Dimargaritales)]
[Dissophora — see Mortierellaceae (Mortierellales)]
[Echinosporangium — a synonym of Lobosporangium: see Mortierellaceae (Mucorales)].
[Ellisomyces — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Fennellomyces — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Endogone — see Endogonaceae (Endogonales])
[Gilbertella — see Choanephoraceae (Mucorales)].
[Gongronella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Halteromyces — see Absidiaceae (Mucorales)].
[Helicocephalum — see Helicocephalidaceae (Zoopagales)]
[Hesseltinella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Hyphomucor — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Kickxella — see Kickxellaceae (Kickxellales)]
[Kirkomyces — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Lentamyces — see Absidiaceae (Mucorales)].
[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)]
[Mycocladus — see Mycocladaceae (Mucorales): a taxon probably based on a mixture of two species, nomen. confus., probably Lentamyces parricida and another mucoralean fungus; see Hoffmann et al., 2009. Citation: Mycocladus Beauverie, 1900 (Ann. Univ. Lyon, Sér. 2, Sciences, Médicine 3: 162); 1 sp. (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 — see Mycotyphaceae (Mucorales)].
[Parasitella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Phascolomyces — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Phycomyces — see Phycomycetaceae (Mucorales)].
[Pilobolus — see Pilobolaceae (Mucorales)].
[Piptocephalis — see Piptocephalidaceae (Zoopagales)]
[Poitrasia — see Choanephoraceae (Mucorales)].
[Protomycocladus — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Radiomyces — see Radiomycetaceae (Mucorales)].
[Rhizomucor — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Rhizopodopsis — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Rhizopus — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Rhopalomyces — see Helicocephalidaceae (Zoopagales)]
[Saksenaea — see Saksenaeaceae (Mucorales)].
[Sigmoideomyces — see Cunninghamellaceae (Mucorales)].
[Spinalia — see Dimagaritaceae (Dimargaritales)]
[Spinellus — see Phycomycetaceae (Mucorales)].
[Spirodactylon — see Kickxellaceae (Kickxellales)]
[Spiromyces — see Kickxellaceae (Kickxellales)]
[Sporodinia — a synonym of Syzygites: see See Synopsis of Genera with Unknown Affinities (Mucorales)].
[Sporodiniella — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Syncephalastrum — see Syncephalastraceae (Mucoirales)].
[Syncephalis — see Piptocephalidaceae (Zoopagales)]
[Syzygites — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Thamnocephalis — see Cunninghamellaceae (Mucorales)].
[Thamnostylum — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Thermomucor — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Tieghemiomyces — see Dimagaritaceae (Dimargaritales)]
[Umbelopsis — see Umbelopsidaceae (Mucorales)]
[Utharomyces — see Pilobolaceae (Mucorales)].
[Zychaea — see Synopsis of Genera with Unknown Affinities (Mucorales)].
[Zygorhynchus — see Dicranophoraceae (Mucorales)].
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Updated Jun 09, 2010