About the Author

This website was based on chapter written by Gerald Benny and R.K. Benjamin in 1991. The chapter was to be included in a book that was never published.

The information has since been updated. If you have comments on or corrections for the website, please email the author .

About the Author

Gerald Benny has studied the Zygomycetes for 42 years. First as a student of Dr. R.K. Benjamin at the Rancho Santa Ana Botanic Garden (Claremont, California) and later in the Department of Botany and more recently, the Department of Plant Pathology, University of Florida, Gainesville, Florida. His expertise is in those members of the Zygomycetes that can be grown in culture [Dimargaritales, Kickxellales, Mortierellales, Mucorales, Zoopagales (Piptocephalidaceae)]. Dr. Benny monographed the Thamnidiaceae (Mucorales) for his Ph.D. dissertation (Benny, 1973) and the results of the study have been published in various journals (Benny, 1992, 1995b, 1995c; Benny and Benjamin, 1975, 1976, 1993; Benny and Schipper, 1992). Other studies of the Mucorales have included those on Mycotyphaceae (Benny et al., 1985), Gilbertellaceae (Benny, 1991), Radiomycetaceae (Benny and Benjamin, 1991; Benny and Khan, 1988; Benny and Samson, 1989), and Utharomyces (Kirk and Benny, 1980). Sigmoideomycetaceae (Zoopagales) was also monographed (Benny et al., 1992) based mostly on herbarium specimens. One paper “Classical morphology in zygomycete taxonomy” (Benny, 1995a) was based on a talk given at the Fifth International Mycological Congress, August 1994, Vancouver, British Columbia, Canada. A chapter was written on the classification and phylogeny of the Trichomycetes and Zygomycetes (Benny and White, 2001). Separate chapters were published on the Trichomycetes and the Zygomycetes (Benny, 2001; Benny et al., 2001). The Zygomycota have been reviewed (Benny 1982, 2012). Echinosporangium was renamed and Mortierella subgenus Gamiella was elevated to generic status (Benny and Blackwell, 2004). Other papers have been on Syncephalis (Ho and Benny, 2007, 2008; Santiago et al., 2011), and the methods used to isolate, culture, preserve and study culturable Zygomycetes (Krug et al., 2004; Benny, 2008). The description of three subphyla (Mucoromycotina, Kickxellomycotina, Zoopagomycotina) of the zygomycetes was included in a multi-authored paper on the phylogenetic classification of the Fungi (Hibbett et al., 2007).


Benny, G. L. 1973. A Taxonomic Revision of the Thamnidiaceae (Mucorales). Dissertation, Claremont Graduate School, Claremont, California. 252 p.

Benny, G.L. 1982. Zygomycetes, pp. 184-195. In S. P. Parker (Ed.). Synopsis and Classification of Living Organisms. Vol. I. McGraw-Hill Book Company, Inc., New York.

Benny, G.L. 1991. Gilbertellaceae, a new family of the Mucorales (Zygomycetes). Mycologia 83: 150-157.

Benny, G.L. 1992. Observations on Thamnidiaceae (Mucorales). V. Thamnidium. Mycologia 84: 834-842.

Benny, G.L. 1995a. Classical morphology in zygomycete taxonomy. Canadian Journal of Botany 73(Suppl. 1): S725-S730.

Benny, G.L. 1995b. Kirkomyces, a new name for Kirkia Benny. Mycologia 87: 922.

Benny, G.L. 1995c. Observations on Thamnidiaceae. VII. Helicostylum and a new genus Kirkia. Mycologia 87: 253-264.

Benny, G.L. 2001. The Zygomycota: Trichomycetes, pp. 147-160. In: D.J. McLaughlin, E.G. McLaughlin, and P.A. Lemke (eds.). The Mycota. Vol. 7A. Systematics and Evolution. Springer-Verlag, Berlin, Germany.

Benny, G.L. 2008. The methods used by Dr. R.K. Benjamin, and other mycologists, to isolate Zygomycetes. Aliso 26: 37-61.

Benny, G.L. 2012. Current systematics of Zygomycota with a brief review of their biology, pp. 55-105. In: J.K. Misra, P. Tewari, and K. Deshmukh (eds.) Systematics and Evolution of Fungi. Science Publishers, Enfield, New Hampshire.

Benny, G.L., and R. K. Benjamin. 1975. Observations on Thamnidiaceae (Mucorales). New taxa, new combinations, and notes on selected species. Aliso 8: 301-351.

Benny, G.L., and R.K. Benjamin. 1976. Observations on Thamnidiaceae (Mucorales). II. Chaetocladium, Cokeromyces, Mycotypha, and Phascolomyces. Aliso 8: 391-424.

Benny, G.L., and R.K. Benjamin. 1991 [1992]. The Radiomycetaceae (Mucorales; Zygomycetes). III. A new species of Radiomyces, and cladistic analysis and taxonomy of the family; with a discussion of the evolutionary ordinal relationships in Zygomycotina. Mycologia 83: 713-735.

Benny, G.L., and R.K. Benjamin. 1993. Observations on Thamnidiaceae (Mucorales). VI. Two new species of Dichotomocladium and the zygospores of D. hesseltinei (Chaetocladiaceae). Mycologia 85: 660-671.

Benny, G.L., R.K. Benjamin, and P. M. Kirk. 1992. A reevaluation of Cunninghamellaceae (Mucorales). Sigmoideomycetaceae fam. nov. and Reticulocephalis gen. nov.; cladistic analysis and description of two new species. Mycologia 84: 615-641.

Benny, G.L., and M. Blackwell. 2004. Lobosporangium, a new name for Echinosporangium Malloch, and Gamsiella, a new genus for Mortierella multidivaricata. Mycologia 96:143-149.

Benny, G.L., R.A. Humber, and J.B. Morton. 2001. The Zygomycota: Zygomycetes, pp. 113-146. In: D.J. McLaughlin, E.G. McLaughlin, and P.A. Lemke (eds.). The Mycota. Vol. 7A. Systematics and Evolution. Springer-Verlag, Berlin, Germany.

Benny, G.L., and S.R. Khan. 1988. The Radiomycetaceae (Mucorales; Zygomycetes). Calcium oxalate crystals on the sporangiolar wall and aerial hyphae. Scanning Microscopy 2(2):1199-1206.

Benny, G.L., P. M. Kirk, and R.A. Samson. 1985. Observations on Thamnidiaceae (Mucorales). III. Mycotyphaceae fam. nov. and a re-evaluation of Mycotypha sensu Benny & Benjamin illustrated by two new species. Mycotaxon 22: 119-148.

Benny, G.L., and R.A. Samson. 1989. The Radiomycetaceae (Mucorales, Zygomycetes). II. Ultrastructural morphology of the mature zygospore and comparative morphology of the sporangiola and sporangiolar appendages. Memiors of the New York Botanical Garden 49:11-19.

Benny, G.L., and M.A.A. Schipper. 1992. Observations on Thamnidiaceae (Mucorales). IV. Pirella. Mycologia 84: 52-63.

Benny, G.L., and M.M. White. 2001. The classification and phylogeny of Trichomycetes and Zygomycetes, pp. 39-53. In: J.K. Misra and B.W. Horn (eds.). Trichomycetes and other fungal groups. Science Publishers, Inc., Enfield, New Hampshire, U.S.A.

Hibbett, D.S., M. Binder, J.F. Bischoff, M. Blackwell, P.F. Cannon, O.E. Eriksson, S. Huhndorf, T. James, P.M. Kirk, R. Lücking, H.T. Lumbsch, F. Lutzoni, P.B. Matheny, D.J. McLaughlin, M.J. Powell, S. Redhead, C.L. Schoch, J.W. Spatafora, J.A. Stalpers, R. Vilgalys, M.C. Aime, A. Aptroot, R. Bauer, D. Begerow, G.L. Benny, L.A. Castelbury, P.W. Crous, Y.C. Dai, W. Gams, D.M. Geiser, G.W. Griffith, C. Gueidan, D.L. Hawksworth, G. Hestmark, K. Kosaka, R.A. Humber, K.D. Hyde, J.E. Ironside, U. Kõljalg, C.P. Kurtzman, K.-H. Larsson, R. Lichtwardt, J. Longcore, J. Miądlikowska, A. Miller, J.-M. Moncalvo, S. Mozley-Standridge, F. Oberwinkler, E. Parmasto, V. Reeb, J.D. Rogers, C. Roux, L. Ryvarden, J.P. Sampaio, A. Schüßler, J. Sugiyama, R.G. Thorn, L. Tibell, W.A. Untereiner, C. Walker, Z. Wang, A. Weir, M. Weiss, M.M. White, K. Winka, Y.-J. Yao and N. Zhang. 2007. A higher-level classification of the fungi. Mycological Research 111: 509-547.

Ho, H.-M. and G.L. Benny. 2007. Two new species of Syncephalis from Taiwan, with a key to the Syncephalis species found in Taiwan. Botanical Studies 48: 319-324.

Ho, H.-M. and G.L. Benny. 2008. A new species of Syncephalis from Taiwan. Botanical Studies 49: 45-48.

Kirk, P.M., and G. L. Benny. 1980. The genus Utharomyces Boedijn (Pilobolaceae: Zygomycetes). Trans. Brit. Mycol. Soc. 75:123-131.

Krug, J.C., G.L. Benny and H.W. Keller. 2004. Coprophilous fungi, pp. 467-499. In: G.M. Mueller, G.F. Bills and M.S. Foster. [eds.]. Biodiversity of fungi: Inventory and monitoring methods. Elsevier Academic Press, Amsterdam, The Netherlands.

Santiago, A.L., G.L. Benny, and L.C. Maia. 2011. Syncephalis aggregata, a new species from the semiarid region of Brazil. Mycologia 103:135-138.

Updated Dec 30, 2011