Classically the Pilobolaceae included two genera, Pilobolus Tode: Fr. and Pilaira van Tieghem (Grove, 1934). These genera have been placed in a subfamily of the Mucoraceae (Fischer, 1892; Schröter, 1893) or in a family of their own (van Tieghem, 1878; Lendner, 1908; Fitzpatrick, 1930; Hesseltine, 1955; Benjamin, 1959, 1979). Utharomyces Boedijn (1958) was added after its description (Hesseltine and Ellis, 1973; Benjamin, 1979). A few authors have included members of the Pilobolaceae in the Mucoraceae (Schröter, 1889; Zycha et al., 1969). Characteristics of Pilobolus, Pilaira and Utharomyces were reviewed by Kirk and Benny (1980) in their definitive study of U. epallocaulus.

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) used by O’Donnell et al. (2001) showed that the Pilobolus — Utharomyces clade (Pilobolaceae) had 93% 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 (98% BP) and strict concensus (95% BP) yielded support for the Pilobolaceae. Based on molecular phylogenetic studies, Pilaira is a member of the Mucoraceae (O’Donnell et al., 2001; Voigt and Olsson, 2008).

Page (1956, 1962) studied the effect of light on the development of the trophocyst and the subsequent production of sporangia in Pilobolus. Sporangia of Pilobolus spp. aid in the dispersal of lungworm disease larvae (Dictyocaulus viviparous) in cattle (Bos primigenus L.) and elk [Cervus canadensis (Erxieleben)] (Eysker, 1991; Foos, 1997; Panuska, 2006). Kock et al. (2001) reported the presence of 3-hydroxyoxylipins in the sporangia, on the columella and the developing sporangiospores, of Pilobolus. Sporangial discharge has been studied and illustrated, and a video supplied of the process, by Yafetto et al. (2008). Utharomyces does not appear to have any economic value.

Pilaira, Pilobolus and Utharomyces are included in the Pilobolaceae by Cannon and Kirk (2007) whereas the family is reduced to the latter two genera by Kirk et al. (2008). Pilobolaceae are primarily coprophilous although there are a few reports of their having been isolated from soil (Grove, 1934; Kirk and Benny, 1980).


Pilobolaceae Corda ex Corda, 1842 (Anleit. Stud. Mycol., p. 71).
= Pilobolaceae Corda, 1837 (Icon. Fung. 1: 22).

Subaerial mycelium sparse, coenocytic when young, branched, producing trophocysts, which give rise to one, or rarely two or more sporophores. Sporophores usually unbranched. Sporangia multispored, columellate, wall persistent, dark, cuticularized, often covered with crystals, with a conspicuous subsporangial vesicle, forcibly discharged or passively detached, with or without the columella and the upper part of the sporophore. Suspensors apposed, often entwined, without appendages. Zygospores brown to black, formed within or at the surface of the substrate; zygosporangium hyaline, rough or wavy.

Type genus: Pilobolus

Key to the Genera of Pilobolaceae

A. Subsporangial vesicle attached to the base of the sporangium; dehiscence of the sporangium by a buildup of turgor pressure in the vesicle inducing rupture of circumscissile zone of weakness at the base of the sporangial wall; sporangium actively shot away up to 2 meters from the sporangiophore; hemin or dung extract required for growth and sporulation —— Pilobolus

AA. Subsporangial vesicle separated from the sporangium by a short stalk; sporangium dehisced as a result of the elongation of the sporangiophore, the sporangial wall breaks after contacting a solid surface; sporangium not shot away; hemin or dung extract not required for growth and sporulation —— Utharomyces

Synopsis of Genera

PILOBOLUS Tode: Fr., 1832 (Syst. Mycol. 3: 312); 10 to 13 spp. + 2 varieties (Coemans, 1861; Klein, 1872; Brefeld, 1881; Grove, 1934 -ILLUS. and KEY TO SPP.; McVickar, 1942 -illus.; Lyr, 1954 -illus.; Hu et al., 1989 -revision; Yafetto et al., 2008-sporangial discharge)
= Pilobolus Tode, 1784 (In Schrift. Berl. Gesell. naturf. Fr., 5: 46).
= Pycnopodium Corda, 1842 (Icon. Fung. 5: 18; fide Hesseltine, 1955).

UTHAROMYCES Boedijn, 1959 (Sydowia 12: 340); 1 sp. (Boedijn, 1958; Kirk and Benny, 1980 -ILLUS.).

Genus Not Accepted Here but Included in Pilobolaceae by Some Authors

[Pilaira — see Mucoraceae (Mucorales)].


Benjamin, R.K. 1959. The merosporangiferous Mucorales. Aliso 4: 321-433.

Benjamin, R.K. 1979. Zygomycetes and their spores, pp. 573-616. In: B. Kendrick (Ed.). The whole fungus: the sexual—asexual synthesis. Vol. 2. National Museums of Canada, Ottawa, Canada.

Boedijn, K. B. 1959 [1958]. Notes on the Mucorales of Indonesia. Sydowia 12: 321-362.

Brefeld, O. 1881. Pilobolius, pp. 60-80. In: O. Brefeld. Botanische Untersuchungen über Schimmelpilze. Leipzig, Germany, Verlag von Arthur Felix.

Cannon, P.F., and P.M. Kirk. 2007. Fungal families of the world. CAB International. Wallingford, United Kingdom. 456 p.

Coemans, W. 1861. Monographie du genre Pilobolus, Tode, spécialement étudié au point de vue anatomie et physiologique. Mémoires Couronnés et Mémoires des Sevantes Étrangers, Académie Royale des Science, des Lettres et des Beaux-Arts de Belgique 30: 3-68.

Eysker, M. 1991. Direct measurement of dispersal of Dictyocaulus viviparous in sporangia of Pilobolus species. Research in Veterinary Science 50:29-32.

Fischer, A. 1892. Phycomycetes In: L. Rabenhorst. Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz. Zweite Auflage. Band I, Abth. IV. Eduard Kummer, Leipzig, Germany. 505 p.

Fitzpatrick, H.M. 1930. The Lower Fungi. Phycomycetes. McGraw-Hill Book Co., Inc., New York. U.S.A. 331 p.

Foos, K.M. 1997. Pilobolus and lungworm disease affecting elk in Yellowstone National Park. Mycological Research 101:1535-1536.

Grove, W. B. 1934. A systematic account and arrangement of the Pilobolidae, pp. 190-224, In A. H. R. Buller. Researches on Fungi. Vol. 6. London, Longmans Green and Co. 513 p.

Hesseltine, C.W. 1955. Genera of Mucorales with notes on their synonymy. Mycologia 47:344-363.

Hesseltine, C.W., and J.J. Ellis. 1973. Mucorales, pp. 187-217, In G. C. Ainsworth, F. K. Sparrow, and A. S. Sussman (Eds.). The Fungi. Vol. 4b. Academic Press, New York. 504 p

Hu, F.-m., R.-y. Zheng, and G.-q. Chen. 1989. A redelimitation of the species of Pilobolus. Mycosystema 2:111-133.

Kirk, P.M., and G. L. Benny. 1980. The genus Utharomyces Boedijn (Pilobolaceae: Zygomycetes). Transactions of the British Mycolological Society 75:123-131.

Kirk, P.M., P.F. Cannon, D.W. Minter, and J.A. Stalpers. 2008. Ainsworth & Bisby’s Dictionary of the Fungi. 10th Ed. Wallingford, United Kingdom, CAB International. 771 p.

Klein, J. 1872. Zur Kenntniss des Pilobolus. Jahrbücher für wissenschaftliche Botanik 8:305-381.

Kock, J.L.F., T. Strauss, C.H. Pohl, D.P. Smith, P.J. Botes, E.E. Pretorius, T. Tepeny, O.Sebolai, A. Botha, and S. Nigam. 2001. Bioprospecting for novel oxylipins in Fungi: the presence of 3-hydroxyoxylipins in Pilobolus. Antonie van Leeuwenhoek 80:93-99.

Lendner, A. 1908. Les Mucorinées de la Suisse. Beiträge zur Kryptogamenflora der Schweiz. III. K.-J. Wyss, Bern, Switzerland. 180 p.

Lyr, H. 1954. Zur Kenntnis der Sexualverhältnisse der Gattung Pilobolus. Archiv für Protistenkunde 99:252-293.

McVickar, D.L. 1942. The light-controlled diurnal rhythm of asexual reproduction in Pilobolus. American Journal of Botany 29:372-380.

O’Donnell, K., F.M. Lutzoni, T.J. Ward, and G.L. Benny. 2001. Evolutionary relationships among mucoralean fungi (Zygomycota): Evidence for family polyphyly on a large scale. Mycologia 93: 286-296.

Page, R.M. 1956. Studies on the development of asexual reproductive structures in Pilobolus. Mycologia 48:206-224.

Page, R.M. 1962. Light and the asexual reproduction of Pilobolus. Science 138:1238-1245.

Panuska, C. 2006. Lungworms of ruminants. Veterinary Clinics of North America: Food Animal Practice 22:583-593.

Schröter, J. 1889. VIII. Ordn. Zygomycetes. Brefeld, pp. 197-225. In: Dr. F. Cohn’s Kryptogamen-Flora von Schlesien. Band IIIa. Pilze. J.U. Kern’s Verlag (Max Müller), Breslau, Germany.

Schröter, J. 1893. Mucorineae, pp. 119-134. In A. Engler and K. Prantl, Die natürlichen Pflanzenfamilien. I Teil, 1. Abth. Wilhelm Engelmann, Leipzig.

van Tieghem, P. 1878. Troisième mémoire sur les Mucorinées. Annales des Sciences Naturelles, Botanique, Séries VI, 4:312-399.

Voigt, K., and I. Olsson. 2008. Molecular phylogenetic and scanning electron microscopical analysis places the Choanephoraceae and the Gilbertellaceae in a monophyletic group within the Mucorales (Zygomycetes, Fungi). Acta Biologica Hungarica 59: 365-383.

Yafetto, L., L.Carroll, Y. Cui, D.J. Davis, M.W. Fischer, A.C. Henterly, J.D. Kessler, H.A. Kilroy, J.B. Shidler, J.L. Stolze-Rybczynski, Z. Sugawara, N.P. Money. 2008. The fastest flights in nature: high-speed spore discharge mechanisms among fungi. PLOS One 3:e3237, 5 p.

Zycha, H., R. Siepmann, and G. Linnemann. 1969. Mucorales eine Beschreibung aller Gattungen und Arten dieser Pilzgruppe. Lehre, J. Cramer. 355 p.

Updated Jun 06, 2010