Belongs within: Psathyrellaceae.
Coprinellus is a genus of lignicolous, terrestrial or coprophilous species of coprinoid mushrooms.
Sinister ink?
Published 3 July 2024
With the recent arrival of winter rains (after a record-breaking long and dry summer that seemed to completely bypass autumn), we have reached mushroom season here in Perth. Among the commonly seen varieties of mushrooms in this region are the ink caps, so-named from the way in which their gills dissolve into puddles of black goo as they release their spores. Until recently, ink caps were united as species of the genus Coprinus but their resolution as polyphyletic by molecular data lead to the genus’ own dissolution by Redhead et al. (2001). Among the larger of the segregate taxa to emerge from this division was the genus Coprinellus.
Over forty species were included in Coprinellus by Redhead et al. (2001) and that number has continued to increase as subsequent authors have described new species or assigned further ex-Coprinus as their relations became resolved. Though confidently united by molecular data, the species of Coprinellus are diverse in structure. Most have gills that are at least somewhat deliquescent, in typical ink cap fashion, though a few such as C. disseminatus do not. The developing mushroom bears a protective veil or a covering of minute hairs (setules); a few species have some combination of both. The cap may or may not hang down around the sides, and basidia are polymorphic. Species may grow from wood, soil or animal dung.
Species of Coprinellus have been divided between three sections, largely on the basis of their external covering (Schafer 2010). Members of section Setulosi bear a covering of setules on the cap and stem that is usually easily seen with a hand lens. Section Domestici is characterised by a veil of floccose scales. In section Micacei, the veil is composed of glistening granules that may be washed off and overlooked in mature specimens. Molecular phylogenetic analysis of Coprinellus by Nagy et al. (2012b) identified major clades within the genus that largely lined up with the pre-existing sections though a handful of species combining setules and veils were scattered among the Domestici and Micacei clades. Nagy et al. (2012a) suggested on the basis of molecular dating that the diversification of setulose Coprinellus may have correlated with the increase in open grasslands during the Miocene. The corresponding increase in large grazing mammals may have provided increased opportunities for dung-growing species, and the covering of hairs has been suggested to provide protection from challenges such as insect browsing or water loss (though I am not sure how they are supposed to ward off insects).
Mushrooms may be the feature of fungi that tends to attract the most immediate attention but they are, of course, only the sexually-produced fruiting bodies of a less readily visible hyphal mass. In some Coprinellus species, the hyphae become dense and colourful prior to the development of the fruiting bodies, sometimes producing a distinctly visible mat. This structure is distinct enough to have historically been given its own generic name, Ozonium. And though Ozonium has been abandoned as a genus owing to the impossibility of identifying individual species (perhaps fortunately, as Ozonium pre-dates Coprinellus in the priority stakes), ‘ozonium’ persists as a term for the structure involved. Coprinellus has also been linked to the anamorphic (asexually sporulating) genus Hormographiella with C. domesticus identified as the teleomorphic form of the anamorphic H. verticillata (Cáceres et al. 2006). Hormographiella species are more diverse in their habits than their associated teleomorphs and may even act as opportunistic pathogens. Cáceres et al. (2006) isolated H. verticillata from skin lesions of patients in Panama, and it had previously been identified in clinical settings such as a cerebro-spinal fluid catheter in France. Hormographiella aspergillata, which has been identified as the anamorphic form of Coprinopsis cinerea, has even been implicated in a fatal pulmonary infection in the Netherlands. Such infections appear to be rare, but they certainly add an extra layer of menace to the idea of handling ink caps.
Systematics of Coprinellus
Characters (from Redhead et al. 2001): Veil (usually of globular cells) present and/or bearing superficial lageniform to bulbous-based filament-bearing cells (setulae) converted into secretory round-tipped pileocystidia; lamellae and pileus fully, partially, or non-deliquescent during sporulation, if non-deliquescent some have geotropic lamellae, otherwise lamellae ageotropic; pleurocystidia present or absent, sometimes concentrated towards margins; pileus membranous or fleshy or very fragile; basidia di-, tri-, or tetramorphic. Ozonium present or absent.
Coprinellus Karsten 1879 (see below for synonymy)RV01
|--*C. deliquescens (Bulliard: Fries) Karsten 1879 (see below for synonymy)RV01
|--C. amphithallus (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. angulatus (Peck) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus angulatus Peck 1873]RV01
|--C. aokii (Hongo) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus aokii Hongo 1966]RV01
|--C. aureogranulatus (Uljé & Aptroot) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. bisporiger (Orton) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 1976 (see below for synonymy)RV01
|--C. bisporus (Lange) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus bisporus Lange 1915]RV01
|--C. brevisetulosus (Arnolds) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. callinus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. congregatus (Bulliard: Fries) Karsten 1879RV01
|--C. curtus (Kalchbrenner) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. digitalis (Batsch) Karsten 1879RV01
|--C. dilectus (Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus dilectus Fries 1838]RV01
|--C. disseminatus (Persoon: Fries) Lange 1938 (see below for synonymy)RV01
|--C. domesticus (Bolton: Fries) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. ellisii (Orton) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus ellisii Orton 1960]RV01
|--C. ephemerus (Bulliard: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. eurysporus (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. fallax (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. fimbriatus (Berkeley & Broome) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. flocculosus (De Candolle) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. furfurellus (Berkeley & Broome) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. hemerobius (Fries) Karsten 1879RV01
|--C. heptemerus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. heterosetulosus (Watling) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. heterothrix (Kühner) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. hiascens (Fries: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. impatiens (Fries: Fries) Lange 1938RV01
|--C. marculentus (Britzelmayr) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. micaceus (Bulliard: Fries) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. pellucidus (Karsten) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. phyllophilus (Karsten) Karsten 1879RV01
|--C. plagioporus (Romagnesi) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. proximellus (Karsten) Karsten 1879RV01
|--C. pyrrhanthes (Romagnesi) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. radians (Desmazières) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. rapidus (Fries) Karsten 1879RV01
|--C. sassii (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. sceptrum (Jungh.) Karsten 1879RV01
|--C. schroeteri (Karsten) Karsten 1879RV01
|--C. sclerocystidiosus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. singularis (Uljé) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus singularis Uljé 1988]RV01
|--C. sororius Karsten 1881RV01
|--C. subdisseminatus (Lange) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. subimpatiens (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. subpurpureus (Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. tardus (Karsten) Karsten 1879RV01
|--‘Coprinus’ tomentosus [=C. lagopus var. tomentosus, *Lentispora tomentosa]RV01
|--C. truncorum (Scopoli: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--C. velaris (Fries) Karsten 1879RV01
|--C. velatopruinatus (Bender) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
|--‘Pseudocoprinus’ venustus McKnight & Allison 1969RV01
|--C. verrucispermus (Josserand & Enderle) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
`--C. xanthothrix (Romagnesi) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 (see below for synonymy)RV01
Coprinellus Karsten 1879 [incl. Coprinusella (Peck) Zerov 1979 (nom. inv.), Ephemerocybe Fayod 1889, Lentispora Fayod 1889, Pseudocoprinus Kühner 1928]RV01
Coprinellus amphithallus (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus amphithallus Lange & Smith 1953]RV01
Coprinellus aureogranulatus (Uljé & Aptroot) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus aureogranulatus Uljé & Aptroot in Uljé, Aptroot & van Iperen 1998]RV01
Coprinellus bisporiger (Orton) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 1976 [=Coprinus bisporiger Orton 1976]RV01
Coprinellus brevisetulosus (Arnolds) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus brevisetulosus Arnolds 1982]RV01
Coprinellus callinus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus callinus Lange & Smith 1953]RV01
Coprinellus curtus (Kalchbrenner) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus curtus Kalchbrenner in Thümen 1876]RV01
*Coprinellus deliquescens (Bulliard: Fries) Karsten 1879 [=Agaricus deliquescens; incl. Coprinus silvaticus, Coprinusella silvatica (Peck) Zerov 1979 (nom. inv.)]RV01
Coprinellus disseminatus (Persoon: Fries) Lange 1938 [=Coprinarius disseminarus (Persoon: Fries) Quélet 1886, Coprinellus disseminata, Coprinus disseminatus, Psathyrella disseminatus, *Pseudocoprinus disseminatus (Persoon: Fries) Kühner 1928]RV01
Coprinellus domesticus (Bolton: Fries) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Agaricus domesticus Bolton 1788]RV01
Coprinellus ephemerus (Bulliard: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Agaricus ephemerus Bulliard 1792, Coprinus ephemerus, *Ephemerocybe ephemera]RV01
Coprinellus fallax (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus fallax Lange & Smith 1953]RV01
Coprinellus fimbriatus (Berkeley & Broome) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus fimbriatus Berkeley & Broome 1871]RV01
Coprinellus flocculosus (De Candolle) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Agaricus flocculosus De Candolle 1815]RV01
Coprinellus furfurellus (Berkeley & Broome) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Agaricus furfurellus Berkeley & Broome 1871]RV01
Coprinellus heptemerus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus heptemerus Lange & Smith 1953]RV01
Coprinellus heterosetulosus (Watling) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus heterosetulosus Watling in Orton 1976]RV01
Coprinellus heterothrix (Kühner) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus heterothrix Kühner 1957]RV01
Coprinellus hiascens (Fries: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Agaricus hiascens Fries 1821]RV01
Coprinellus marculentus (Britzelmayr) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus marculentus Britzelmayr 1893]RV01
Coprinellus micaceus (Bulliard: Fries) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Agaricus micaceus Bulliard 1786, Coprinus micaceus]RV01
Coprinellus pellucidus (Karsten) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus pellucidus Karsten 1882]RV01
Coprinellus plagioporus (Romagnesi) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus plagioporus Romagnesi 1941]RV01
Coprinellus pyrrhanthes (Romagnesi) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus pyrrhanthes Romagnesi 1951]RV01
Coprinellus radians (Desmazières) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Agaricus radians Desmazières 1828]RV01
Coprinellus sassii (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus sassii Lange & Smith 1953]RV01
Coprinellus sclerocystidiosus (Lange & Smith) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus sclerocystidiosus Lange & Smith 1953]RV01
Coprinellus subdisseminatus (Lange) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus subdisseminatus Lange in Lange & Smith 1953]RV01
Coprinellus subimpatiens (Lange & Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus subimpatiens Lange & Smith 1953]RV01
Coprinellus subpurpureus (Smith) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus subpurpureus Smith 1948]RV01
Coprinellus truncorum (Scopoli: Fries) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Agaricus truncorum Scopoli 1772]RV01
Coprinellus velatopruinatus (Bender) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus velatopruinatus Bender 1989]RV01
Coprinellus verrucispermus (Josserand & Enderle) Redhead, Vilgalys & Moncalvo in Redhead, Vilgalys et al. 2001 [=Coprinus verrucispermus Josserand & Enderle in Bender & Enderle 1988]RV01
Coprinellus xanthothrix (Romagnesi) Vilgalys, Hopple & Johnson in Redhead, Vilgalys et al. 2001 [=Coprinus xanthothrix Romagnesi 1941]RV01
*Type species of generic name indicated
References
Cáceres, O., R. Kirschner, M. Piepenbring, H. Schöfer & J. Gené. 2006. Hormographiella verticillata and an Ozonium stage as anamorphs of Coprinellus domesticus. Antonie van Leeuwenhoek 89: 79–90.
Nagy, L. G., J. Házi, B. Szappanos, S. Kocsubé, B. Bálint, G. Rákhely, C. Vágvölgyi & T. Papp. 2012a. The evolution of defense mechanisms correlate with the explosive diversification of autodigesting Coprinellus mushrooms (Agaricales, Fungi). Systematic Biology 61 (4): 595–607.
Nagy, L. G., J. Házi, C. Vágvölgyi & T. Papp. 2012b. Phylogeny and species delimitation in the genus Coprinellus with special emphasis on the haired species. Mycologia 104 (1): 254–275.
[RV01] Redhead, S. A., R. Vilgalys, J.-M. Moncalvo, J. Johnson & J. S Hopple Jr. 2001. Coprinus Pers. and the disposition of Coprinus species sensu lato. Taxon 50: 203–241.
Schafer, D. J. 2010. Keys to sections of Parasola, Coprinellus, Coprinopsis and Coprinus in Britain. Field Mycology 11 (2): 44–51.