PIPTOCEPHALIS de Bary, 1865 (Abhandlungen herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 5: 356); 25 species (de Bary, 1865; Benjamin, 1959, 1966).
= Mucoricola Nieuwland, 1916 (Amer. Midl. Naturalist 4: 82; fide Benjamin, 1959).

Most species haustorial parasites of Mucorales, but one can use ascomycete anamorphs as the host. Sporangiophores dichotomously branched several times; head cells, if formed, and merosporangia produced on the ultimate branches. Merosporangia uni- or multispored, usually cylindrical. Spores variously shaped, when mature released dry or in a spore droplet. Zygospores, when formed, more or less globose; suspensors apposed.

Type species: P. freseniana

Species of Piptocephalis:
P. arrhiza van Tieghem & Le Monnier, 1873 (Annales des Sciences Naturelles, Botanique, Séries V, 17:366) (Benjamin, 1966; Gräfenhan, 1999; Indoh, 1962; Mangin, 1899; Naumov, 1954; van Tieghem and Le Monnier, 1873).
P. benjaminii (Embree) R.K. Benjamin, 1963 (Aliso 5:284) (Benjamin, 1963, 1966; Gräfenhan, 1998; Young, 1969).
= Chaetocladium benjaminii, Embree, 1962 (Mycologia 54:305) (Embree, 1962).
P. brijmohanii Mukerji, 1968 (Mycologia 60:326) (Gräfenhan, 1998; Mukerji, 1968).
[[P. corymbifera Vuillemin; synonym of Piptocephalis freseniana (q.v.) fide Gräfenhan, (1998)]
P. cruciata van Tieghem, 1875 (Ann. Sci. Nat., Bot, Séries VI, 1:149) (Christenberry, 1940; Gräfenhan, 1998; Naumov, 1954; Spinedi and Arambarri, 1986; van Tieghem, 1875)
P. curvata Baijal & B.S. Mehrotra, 1968 (Zbl. Bakt., Abt. II, 122:181) (Baijal and B.S. Mehrotra, 1968; Gräfenhan, 1998).
P. cylindrospora Bainier, 1882 (Étude sur les Mucorinées, p. 111) non Piptocephalis cylindrospora Bainier, 1883 (Ann. Sci. Nat., Bot., Sér. VI, 15:92) (Bainier, 1882, 1883; Benjamin, 1966; Gräfenhan, 1998; Leadbeater and Mercer, 1956, 1957b; O’Donnell, 1979; Spinedi and Arambarri, 1986).
= Piptocephalis virginiana Leadbeater & Mercer, 1957 (Trans. Brit. Mycol. Soc. 40:461) (Berry, 1959; Berry and Barnett, 1957; Leadbeater and Mercer, 1957a).
P. debaryana B.S. Mehrotra, 1960 (Proceedings of the National Academy of Sciences, India, Section B, 30:371) (Gräfenhan, 1998; B.S. Mehrotra, 1960).
P. dichotoma Krzemieniewska & Badura, 1954 (Acta Societatis Botanicorum Poloniae 23:733) (Krzemieniewska and Badura, 1954).
[[P. digitata McDaniel & Hindal (Mycologia 74:271—[nomen nudum, without a description, illustration, and a Latin dignosis) (Gräfenhan, 1998; McDaniel and Hindal, 1982)].
P. fimbriata Richardson & Leadbeater, 1972 (Trans. Brit. Mycol. Soc. 58:206) (Gräfenhan, 1998; Richardson and Leadbeater, 1972).
P. freseniana De Bary, 1865 (Abh. Senckenb. Natf. Ges. 5:356) (Bainier, 1882; Brefeld, 1872; Christenberry, 1940; De Bary, 1865; Gräfenhan, 1998; Morini, 1905; Vuillemin, 1902).
= Piptocephalis repens van Tieghem and Le Monnier, 1873 (Annales des Sciences Naturelles, Botanique, Séries V, 17:364) (Bainier, 1882; Naumov, 1954; O’Donnell, 1979; Ou, 1940; Trufem, 1984; van Tieghem and Le Monnier, 1873; Viriato and Trufem, 1985).
= Piptocephalis corymbifera Vuillemin, 1887 (Bull. Soc. Mycol. France 3:111) (Vuillemin, 1887 [no ill.])
P. fusispora van Tieghem, 1875 (Ann. Sci. Nat., Bot, Séries VI, 1:146) (Morini, 1905; Naumov, 1954; van Tieghem, 1875).
P. graefenhanii H. M. Ho, 2006 (Botanical Studies 47:453) (Ho, 2006).
P. indica B.S. Mehrotra and Baijal, 1964 (Sydowia 17:171) (Baker, 1979; Benjamin, 1966; Gräfenhan, 1998; Ho, 2003; B.S. Mehrotra and Baijal, 1964).
= ?[Piptocephalis indica var shantinketna B.R. Mehrotra & Kakkar, 1969 (Sydowia 23:200); a possible synonym of Piptocephalis indica fide Gräfenhan (B.R. Mehrotra and Kakkar, 1969).
P. lemonnieriana Vuillemin, 1902 (Bull. Mensuel Séances Soc. Sci. Nancy, Séries 3, 3:47) (Vuillemin, 1902).
P. lepidula (Marchal) R.K. Benjamin, 1959 (Aliso 4:345) (Benjamin, 1959, 1966; Gräfenhan, 1998; Indoh, 1962; McDaniel and Hindal, 1979—host range; Mil’ko, 1974)
= Piptocephalis fusispora var. lepidula Marchal, 1891 (Bull. Soc. Roy. Bot. Belgique 2(30): 136)
[[P. macrospora van Beyma thoe Kingma; = synonym of Piptocephalis tieghemiana fide Gräfenhan, 1998)].
[[P. macrocephala Mirza, Khan, Begum & Shagufta, 1979 ([Mucorales of Pakistan, p. 123—nomen nudum, no Latin diagnosis or illustration)].
P. microcephala van Tieghem, 1875 (Ann. Sci. Nat., Bot, Séries VI, 1:147) (Benjamin, 1966; Gräfenhan, 1998; Naumov, 1954; van Tieghem, 1875).
P. minuta Kuzuha, 1976 (Journal of Japanese Botany 51:123) (Gräfenhan, 1998; Kuzuha, 1976).
P. monospora Mangin, 1899 [Journal de Botanique (Paris) 13:376] (Mangin, 1899).
P. pseudocephalia P.M. Kirk, 1978 (Trans. Brit. Mycol. Soc. 70: 337) (Gräfenhan, 1998; Jeffries, 1979; Kirk, 1978).
[[P. repens van Tieghem and Le Monnier; a synonym of Piptocephalis freseniana fide Gräfenhan (1998)].
[P. richardii Baijal & B.S. Mehrotra ex Gräfenhan, 1998 ([Taxonomic revision of the genus Piptocephalis, p. 59) (Baijal and B.S. Mehrotra, 1968; possibly in B.S. Mehrotra and Baijal, 1964; Gräfenhan, 1998)].
P. sphaerocephala Mangin, 1899 [Journal de Botanique (Paris) 13:376] (Mangin, 1899; Naumov, 1954).
P. sphaerospora van Tieghem, 1875 (Ann. Sci. Nat., Bot, Séries VI, 1:150) (van Tieghem, 1875).
P. tieghemiana Matruchot, 1900 (Bull. Soc. Mycol. France 16:58) (Benjamin, 1966; Gräfenhan, 1998; Matruchot, 1900; Mil’ko, 1974; Mistry, 1977; O’Donnell, 1979).
= Piptocephalis macrospora van Beyma thoe Kingma. 1944 (Antonie van Leeuwenhoek 10:43) fide Gräfenhan (1998) (van Beyma thoe Kingma, 1944).
P. unispora R.K. Benjamin, 1966 (Mycologia 42:23) (Benjamin, 1966; Cuthbert and Jeffries, 1984—host range; Gräfenhan, 1998; Jeffries and Young, 1975a, 1975b, 1976a, 1976b, 1978; Young, 1969).
P. virginiana Leadbeater & Mercer, 1957 (Trans. Brit. Mycol. Soc. 40:461) (Berry, 1959—parasitism; Berry and Barnett, 1957—host range; Leadbeater and Mercer, 1957a, 1957b; Manocha, 1975—spore morphology and biochemistry; Manocha et al., 1997—biochemistry; O’Donnell, 1979; Young, 1969).
P. xenophila Dobbs & English, 1954 (Trans. Brit. Mycol. Soc. 37:375) (Benjamin, 1959; Dobbs and English, 1954; Gräfenhan, 1998; Leadbeater and Mercer, 1957b).

Piptocephalis can be isolated from many habitats. Many taxa were isolated from rodent dung by Dr. R.K. Benjamin. I have isolated a few species from soil but members of Piptocephalis do not appear to be as common in soil as is Syncephalis. Isolation of Piptocephalis can be relatively easy if you have watch makers forceps used by electron microscopists and fine needle such as stainless steel minuten insect pins that can be mounted in inoculating needle handles. Spores from spore droplets can be transferred using either the minuten insect pins or forceps whereas the dry spores should be collected using forceps. Spores are then inoculated on an appropriate host.

Benjamin (1959) recommended using Cokeromyces recurvatus, or for some Piptocephalis species, Umbelopsis ramannianus, as the host for Piptocephalis spp. The host can be grown on a relatively nutrient-rich culture medium, such as MEYE or Emerson’s YpSs agars. Jeffries and Kirk (1976) recommended the use of C. recurvatus yeast cells obtained from liquid culture as the initial host for Piptocephalis, and other merosporangiferous mucorales.

Benjamin (1959, 1963, 1966), and Kirk (1978) described and illustrated four species of Piptocephalis. The genus was monographed by Gräfenhan (1998) as his M.S. research but the final version has not been published.

Baijal, U., and B.S. Mehrotra. 1968. An interesting species of Piptocephalis. Zbl. Bakt, Abt. II, 122:181-184.

Bainier, G. 1882. Étude sur les Mucorinées. Thèse. Paris, F. Pichon et A. Cotillon, Imprimeurs. 136 p.

Bainier, G. 1883. Observations sur les Mucorinées. Ann. Sci. Nat., Bot., Sér. VI, 15:70-104.

Baker, K.K. 1979. Ultrastructural development of merosporangia in the mycoparasite Piptocephalis indica. Can. J. Bot. 57:2460-2465.

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

Benjamin, R.K. 1963. Addenda to “The merosporangiferous Mucorales” II. Aliso 5: 273-288.

Benjamin, R.K. 1966. The merosporangium. Mycologia 58:1-42.

Berry, C.R. 1959. Factors affecting parasitism of Piptocephalis virginiana on other Mucorales. Mycologia 51:824-832.

Berry, C.R., and H.L. Barnett. 1957. Mode of parasitism and host range of Piptocephalis virginiana. Mycologia 49:374-386.

Brefeld, O. 1872. Botanische Untersuchungen über Schimmelpilze. I. Heft: Mucor mucedo, Chaetocladium jones’ii, Piptocephalis freseniana. Zygomyceten. Verlag von Arthur Felix, Leipzig, Germany. 64 p.

Christenberry, G.A. 1940. A taxonomic study of the Mucorales in the southeastern United States. Journal of the Elisha Mitchell Scientific Society 56:333-366.

Cuthbert, A., and P. Jeffries. 1984. Mycoparasitism by Piptocephalis unispora within the Mortierellaceae. Trans. Brit. Mycol. Soc. 83:700-702.

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Ho, H.-M. 2003. The merosporangiferous fungi from Taiwan (III): Three new records of Piptocephalidaceae (Zoopagales, Zygomycetes). Taiwania 48:53-59.

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Indoh, H. 1962. Notes on Japanese Mucorales I. Trans. Mycol. Soc. Japan 3:24-28.

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Jeffries, P., and T.W.K. Young. 1975b. Ultrastructure of the sporangiospore of Piptocephalis unispora (Mucorales). Arch. Mikrobiol. 105:329-333.

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Jeffries, P., and T.W.K. Young. 1976b. Ultrastructure of infection of Cokeromyces recurvatus by Piptocephalis unispora (Mucorales). Arch. Mikrobiol. 109:277-288.

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Manocha, M.S., D. Xiong, and V. Govindsamy. 1997. Isolation and partial characterization of a complementary protein from the mycoparasite Piptocephalis virginiana that specifically binds to two glycoproteins at the host cell surface. Can. J. Microbiol. 43:625-632.

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Updated Nov 06, 2007