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Coprinoid Mushrooms: The Inky Caps  

[ Basidiomycota > Agaricales > Agaricaceae / Psathyrellaceae . . . ]

by Michael Kuo

Inky caps are fascinating mushrooms. They are saprobes, assisting in the decomposition of wood, dung, grassy debris, forest litter, and so on. Most of the species have black spore prints and gills that liquefy, at least partially, as the mushroom matures. The resulting "ink" provides the common name for the inky caps, and can actually be used as writing ink.

But the mushrooms, of course, do not have the production of ink for writing in mind. Rather, liquefying the gills is a clever strategy for dispersing spores more efficiently. The gills liquefy from the bottom up as the spores mature. Thus the cap peels up and away, and the maturing spores are always kept in the best position for catching wind currents. As this happens, the shape of the cap progresses from more or less oval (when seen from the side) to broadly bell-shaped and, eventually, more or less flat as the spores nearest to the stem are exposed to the air currents.

Identification of inky caps ranges from fairly easy (Coprinus comatus and Coprinopsis atramentaria, for example, are common and widely known) to extremely difficult, especially when it comes to the tiny ones. Simply getting some inky caps home to study can be a challenge, since many are so ephemeral that they appear, liquefy, and turn into black goo within a matter of hours. Identification of these short-lived mushrooms (did I mention that they all look pretty much the same?) hinges on microscopic examination of various erudite features, and is an enterprise best left to folks who enjoy such endeavors.

Recent DNA studies have resulted in some fascinating rearrangements of the coprinoid mushrooms. The fact that a mushroom's gills liquefy and turn into black goo, it turns out, does not necessarily mean it is closely related to other mushrooms that do the same thing. Thus the traditional genus Coprinus, which was conceived on the idea that deliquescing gills (and other physical attributes) indicated genetic relationship, turns out to hold mushrooms that are so far apart, genetically, that they do not even belong to the same family, let alone the same genus. For more on "convergent evolution" (the term used when organisms develop similar features independently, without being closely related) see What, If Anything, Is a Gilled Mushroom?"

According to Redhead and collaborators (2001), coprinoid mushrooms must now be distributed among four genera: Coprinus, Parasola, Coprinopsis, and Coprinellus. While the researchers found physical features that, for the moment, appear to allow separation of these genera, the genera are defined by DNA results and we should not expect the correspondence of physical features to be anything more than coincidental—which means that exceptions may arise. The researchers changed genus names for a large number of mushrooms that have not been subjected to DNA analysis (as well as many that have), on the assumption that the apparent morphological separators among mushrooms that have been tested will remain viable when the rest of the former coprinuses are tested.

Coprinus, in its new and more restricted sense, contains only a few species. The shaggy mane, Coprinus comatus, is the best known. The features shared by species of Coprinus but not by other coprinoid mushrooms, it turns out, include the presence of a ring on the stem, the frequently pinkish young gills, and the presence of a string-like strand of fibers in the hollow stem. Species of Coprinus (along with species of Podaxis, Montagnea, and Xerocoprinus, which represent desert adaptations—gnarly, dried-up things that look like Frankenstein's mockery of the shaggy mane) belong in the Agaricaceae family, more closely related to species of Agaricus and Lepiota than to other coprinoid mushrooms, which belong to the Psathyrellaceae family.

The genus Parasola is comprised of mostly tiny, umbrella-like species that have no universal veil and therefore feature caps that lack scales, patches, granules, and so on. Parasola plicatilis is the best known species.

Coprinopsis and Coprinellus are harder to separate without microscopic analysis. Veils are often more noticeable in Coprinopsis, but this is not always the case. Species with an ozonium (an orange mat of fibers around the stem base) belong in Coprinellus, as do species with mica-like granules on the cap surface and species that are only partially deliquescent. Ultimately, however, microscopic features (especially the type of pileipellis and the disposition of veil elements) must be consulted for most species. For a clear summary of apparent morphological separators among coprinoid genera, see the excellent table (Table 2 on page 36) in Keirle and collaborators (2004).

Until an in-depth, species-level DNA study of coprinoid mushrooms is done, most of the species are still defined as they have always been defined—which means using a microscope if you want to identify them successfully. While naked-eye features can narrow down possibilities, especially with some of the larger species, microscopic differences have been used to define most of the coprinoid species. Spore morphology is often important, as is the anatomy of the pileipellis, the hymenium, and the veil elements. I recommend using a KOH mount, stained with something like phloxine, in order to see the features that define coprinoid mushrooms. A Roman aqueduct section provides a good start for microscopic analysis, but you may find you need to "scalp-section" the center portion of young caps in order to find veil elements; slice a very thin strip from the cap surface.


Coprinus comatus

Parasola plicatilis

Coprinoid sp.

Coprinus comatus

Coprinellus disseminatus

Coprinopsis variegata

Coprinellus domesticus

Coprinellus disseminatus

Coprinellus hiascens

© MushroomExpert.Com

Key to 82 North American Coprinoid Taxa

Note: With a few exceptions, I have included only taxa that have been reported from North America in one of the sources listed below the key.

1.Mushroom fairly large (mature cap usually over 5 cm); whitish with brownish scales, at least over the center; oval when young; gills whitish to pinkish when young, turning to black goo with maturity; stem with a ring, hollow when sliced, with a strand of fibers running through the hollow cavity; often growing in grassy urban areas, or in disturbed ground (ditches, roadsides, etc.) but not on dung; hymenial cystidia absent.

1.Not completely as above.

2.Known from the Pacific Northwest; stem 35-50 cm long; spores 17-20 µ long.
Coprinus colosseus

2.Distribution various; stem much shorter than above; spores 8-18 µ long (Coprinus comatus in the wide sense).

3.Spores with a central apical pore; widely distributed in North America.

3.Spores with an eccentric pore; distribution uncertain (varieties originally recorded from western North America).

4.Spores 14-18 µ long; pore very eccentric.
Coprinus comatus
var. excentricus

4.Spores 8-15 µ long; pore slightly eccentric.
Coprinus comatus
var. caprimammillatus

5.Growing on dung or manured soil—sometimes on compost or decaying straw.

5.Growing elsewhere.

6.Ring usually present on stem.

6.Ring absent.

7.Mature cap tiny (usually under 1 cm across); spores shaped like fat lemons.
Coprinus ephemeroides

7.Mature cap larger than above; spores variously shaped. The subsequent species are reminiscent of miniature, dung-loving shaggy manes.

8.Growing on the dung of horses or cows.

8.Growing on the dung of wild animals (deer, rabbits, etc.).

9.Stem to 8 cm high and 4 mm thick; spores 14-18 µ long.
Coprinus umbrinus

9.Stem to 15 cm high and 8 mm thick; spores 17-26 µ long.
Coprinus sterquilinus

10.Mature cap 5-6 cm across; stem white, becoming brownish to black; spores "clear translucent chestnut brown" in KOH; clamp connections present.
Coprinus spadiceisporus

10.Mature cap under 4 cm across; stem white becoming pinkish at the apex; spores nearly black in KOH; clamp connections absent.
Coprinus roseistipitatus

11.Mushroom arising from sclerotia (black knots of tissue up to 1 cm across) that are buried in the, um, substrate; mushroom identifier, having determined this, quite possibly in need of a new hobby.

11.Mushroom not arising from sclerotia.

12.On the dung of cows; recorded from one location in the Netherlands, and from Hawaii; perhaps to be expected in North America.
Coprinopsis sclerotiorum

12.On the dung of sheep; recorded from Montana and from Michigan.
Coprinus sclerotigenus

13.Cap up to 4 cm across, snow white when young, covered with soft mealy granules that are easily wiped off; found primarily on the dung of cows; spores 15-19 µ long.
Coprinopsis nivea

13.Not completely as above.

14.Pileocystidia present.

14.Pileocystidia absent.

15.Velar sphaerocysts also present on cap surface.

15.Velar sphaerocysts absent.

16.Spores more or less six-sided, 9-14 µ long.
Coprinellus marculentus

16.Spores more or less ellipsoid, 13-17 µ long.
Coprinellus heptemerus

17.Basidia 2-spored.

17.Basidia 4-spored.

18.On dung, compost, or straw; pileocystidia thin-walled; pleurocystidia absent; clamp connections absent.
Coprinellus bisporus

18.Usually on dung; some pileocystidia thick-walled; pleurocystidia present; clamp connections present.
Coprinellus sassii

19.Pleurocystidia absent.

19.Pleurocystidia present.

20.Spores 9.5-13 µ long, with an eccentric pore; some pileocystidia with thick walls.
Coprinellus heterosetulosus

20.Spores 6-9.5 µ long, with a central pore; all pileocystidia with thin walls.
Coprinellus pellucidus

21.Clamp connections present.
Coprinellus ephemerus

21.Clamp connections absent.

22.Spores with a central pore.
Coprinellus stellatus

22.Spores with an eccentric pore.
Coprinellus congregatus

23.Spores covered with a conspicuous sheath.
Coprinopsis narcotica

23.Spores lacking a sheath.

24.Spores rounded-triangular to heart-shaped; cap under 1 cm across.
Parasola misera

24.Spores not as above; cap variously sized.

25.Velar elements on cap round or nearly so (scalp-section the disc of young specimens).

25.Velar elements on cap sausage-shaped to tubular or slightly inflated, in chains—but not round.

26.Mature cap 4-10 mm across; veil elements warty.
Coprinopsis stercorea

26.Mature cap 10-25 mm across; veil elements smooth or slightly granular but not warty.
Coprinus patouillardii

27.Mature cap generally more than 15 mm across.

27.Mature cap generally less than 15 mm across.

28.Spores 9.5-11 x 6.5-7.5 µ; stem often rooting; pleurocystidia usually numerous.
Coprinopsis cinerea

28.Spores 11-17 x 7-10 µ; stem not rooting; pleurocystidia sometimes absent or rare.
Coprinopsis macrocephala

29.Spores 13-15 x 7.5-8.5 µ.
Coprinopsis radiata

29.Spores 7.5-10 x 5-5.5 µ.
Coprinopsis pseudoradiata

30.Growing from logs, fallen trees, or large branches—or at the bases of stumps, or from the (usually dead) roots of trees, appearing terrestrial.

30.Growing elsewhere . . . on wood chips; on straw, plant material or sticks; on organic compost; in grass; in disturbed ground; from carpet.

31.Stem with a ring; growing on the wood of alders in western North America; cap to about 2 cm across.
Coprinus alnivorus

31.Stem without a ring; wood preference, range, and other features variable.

32.Mature cap about 1 cm across, or smaller.

32.Mature cap generally larger than 1 cm across.

33.Known from the Pacific Northwest; growing alone on conifer sticks; young cap whitish to grayish under dark gray to black veil; deliquescence only occurring on the edges of gills; pileipellis a cutis; pileocystidia and velar sphaerocysts absent.
Coprinopsis marcida

33.Widely distributed in North America; growing densely gregariously on wood, especially at the bases of stumps or from decaying roots; cap whitish to grayish, without veil; gills not deliquescing; pileipellis an epithelium; pileocystidia and velar sphaerocysts present.

34.Mushrooms arising from a rust colored to orange carpet of fuzz (an ozonium); pileipellis an epithelium.

34.Mushrooms not arising from an ozonium; pileipellis variable.

35.Spores 6-10 µ long; cheilocystidia subglobose, ellipsoid, or broadly utriform.

35.Spores 8.5-12 µ long; cheilocystidia lageniform.
Coprinellus radians

36.Cap prominently scaly.

36.Cap with granules, hairs, tiny scales over the center only, scattered tiny scales—or relatively naked.

37.Scales orangish brown, small and tightly affixed to the cap (reminiscent of scales in Agaricus); stem with a booted appearance, also adorned with orangish brown scales.

37.Scales whitish to yellowish or tan, woolly to cottony and easily scraped off the cap surface; stem usually not with a booted appearance, whitish and smooth.

38.Growing from well decayed logs or stumps in hardwood forests; scales on cap large and patch-like; stem terminating in a rimmed bulb.

38.Growing in urban areas in woodchips or from stumps—or, especially, from the root systems of dying, dead, or recently removed trees, often in lawns; scales small, not patch-like; stem base not a rimmed bulb.

39.Cap rusty brown, without veil; stem under 5 mm thick; pileipellis an epithelium with brownish setae; spores 13-16.5 µ long, smooth.

39.Not completely as above.

40.Cap whitish when young; spores warty.
Coprinopsis insignis

40.Young cap not whitish; spores smooth or warty.

41.Young cap yellow, brownish yellow, honey colored, yellowish, brownish orange, or orangish.

41.Young cap gray to brown or brownish.

42.Veil disposed as mica-like granules on cap surface; velar sphaerocysts present.

42.Veil absent, or tissue-like; velar sphaerocysts absent.

43.Caulocystidia present. (Note: preliminary DNA evidence suggests that Coprinellus micaceus and Coprinellus truncorum may be genetically identical.)

43.Caulocystidia absent.
Coprinellus truncorum

44.Cap surface smooth or, with a hand lens, very finely hairy; spores warty; pileocystidia present.
Coprinus silvaticus

44.Cap surface with tissuelike veil that breaks up into soft scales; spores smooth; pileocystidia absent.
Coprinellus flocculosus

45.Spores broadly elliptical or nearly round (5.6-8.4 x 4.2-5 µ); apparently known from a single 1965 collection under alder in a Washington (state) campground.
Coprinopsis pinguispora

45.Spores more narrowly elliptical than above.

46.Center of cap depressed; apparently known from a single 1972 collection "around a willow tree in a mixed forest" in Washington.
Coprinopsis depressiceps

46.Center of cap not depressed; widely distributed, common, and well documented in North America.

47.Found in arid, desert or semi-desert ecosystems in western North America.

47.Found in other ecosystems.

48.Gills absent or poorly formed.

48.Gills present and "normal."

49.Mature cap disclike, with gill-like plates on its underside; volva present at stem base (often underground).
Montagnea arenarius

49.Mature cap rounded-cylindrical, more or less like a shaggy mane cap; volva absent.

50.Mushroom 15-45 cm high at maturity; stem 1.5-3 cm thick at apex; in North America known from southern California, southern Arizona, and southern Texas (and probably to be expected in Mexico); spores mostly under 10 µ long, with a distinct pore.

50.Mushroom smaller than above (under 20 cm high, with stem up to 1.5 cm thick at apex); distributed throughout western and southwestern North America; spores variously sized, with or without a distinct pore.

51.Most spores over 9 µ long, with a conspicuous pore.

51.Most spores under 7.5 µ long, with an inconspicuous pore or without an apparent pore.

52.Spores ellipsoid, generally under 7 µ long; gleba olive brown to yellowish brown.
Podaxis argentinus

52.Spores subglobose, generally longer than 7 µ long; gleba reddish brown.
Podaxis microporus

53.Cap adorned with one or more large patches of veil.

53.Cap without large patches of veil.

54.Cap with a single, star-shaped, central patch.

54.Cap with one or several patches, but not as above.
Coprinus xerophilus

55.Cap 1-6 cm wide, rusty brown, without veil, at maturity lined from the margin to the rusty brown center; stem under 5 mm thick; pileipellis an epithelium with brownish setae; spores 13-16.5 µ long, smooth.

55.Not as above.

56.Cap up to 7 cm high, dark brown and then black, adorned with large, gorgeous, contrasting whitish scales; stem up to 30 cm long; spores 14-19 x 9.5-13 µ.
Coprinopsis picacea

56.Not as above.

57.Mature cap tiny (generally under 1 cm across); pileipellis a cutis.

57.Mature cap larger than above—or if tiny, pileipellis not a cutis.

58.Spores shaped like corn kernels, measuring 7-9 x 6-7.5 x 5-5.5 µ; known from Montana and Alberta.
Coprinopsis maysoidispora

58.Spores otherwise shaped; distribution various.

59.Veil hyphae with thick (over .5 µ) walls; spores 7.5-9 x 5.5-8 µ.
Coprinopsis friesii

59.Veil hyphae with thin walls; spore measurements various.

60.Spores subglobose, 9-12 x 8-10 x 7.5-9 µ.
Coprinopsis kubickae

60.Spores subglobose or elliptical, smaller than above.

61.Veil hyphae hyaline; spores 5.5-8 x 4-5 µ; North American distribution uncertain, but possibly widely distributed; apparently harmlessly saprobic on rotting stems and leaves of various plants.
Coprinopsis urticicola

61.Veil hyphae yellow-brown; spores 7-9 x 5-6 µ; known from Canada (British Columbia to Ontario); mycelium parasitic as a winter "snow mold" on various plants (follow the link to the right and scroll to the "Coprinus Snow Mold" section for photographs).
Coprinopsis psychromorbida

62.Mature cap medium sized (at least 3 cm across), honey yellow to yellowish, adorned with mica-like granules.

62.Mature cap variously sized, variously colored, lacking mica-like granules.

63.Caulocystidia present. (Note: preliminary DNA evidence suggests that Coprinellus micaceus and Coprinellus truncorum may be genetically identical.)

63.Caulocystidia absent.
Coprinellus truncorum

64.Mushroom appearing like a miniature shaggy mane, with a cap up to 3 cm across and a stem up to 7 mm thick; spores about 9-10 µ long; known from the Pacific Northwest and from Germany.
Coprinus palmeranus

64.Not as above.

65.Cap fairly large (3-10 cm when mature), appearing naked or with small, inconspicuous scales over the center, gray to grayish brown or brown; pileipellis a cutis; hymenial cystidia conspicuous.

65.Not completely as above.

66.Spores broadly elliptical or nearly round (5.6-8.4 x 4.2-5 µ); apparently known from a single 1965 collection under alder in a Washington (state) campground.
Coprinopsis pinguispora

66.Spores more narrowly elliptical than above.

67.Center of cap depressed; apparently known from a single 1972 collection "around a willow tree in a mixed forest" in Washington.
Coprinopsis depressiceps

67.Center of cap not depressed; widely distributed, common, and well documented in North America.

68.Cap surface smooth or grooved, but naked—without (macroscopic) evidence of a veil, even in button stages (though surface may appear very finely hairy or pubescent with a hand lens, before the pileocystidia collapse); pileocystidia present or absent.

68.Cap surface with evidence of a veil, at least when young; pileocystidia always absent.

69.Fresh cap with purplish hues.

69.Purplish hues absent.

70.Spore print black; fairly commonly collected and well documented.
Coprinellus subpurpureus

70.Spore print with reddish hues; possibly not collected since the 1949 type collection (from a lumber yard in Dexter, Michigan).
Coprinellus fallax

71.Pileocystidia absent.

71.Pileocystidia present.

72.Spores warty.
Coprinus silvaticus

72.Spores smooth.

73.Spores mitre-shaped (like a bishop's hat); growing from charcoal in old burn sites.
Coprinellus angulatus

73.Spores more or less elliptical, ovoid, oval, or subglobose; habitat variable.

74.Cheilocystidia mostly subglobose.

74.Cheilocystidia mostly bottle-shaped, fusoid-ventricose, lageniform, etc. (Coprinus expert Kees Uljé writes: "In species with predominantly globose cheilocystidia . . . a few lageniform ones are sometimes present. In some species with mixed globose and lageniform cheilocystidia, the latter are not always abundant. Therefore identification is not easy. Fortunately there are other characters that may help.")

75.Pileocystidia subcapitate; spores 9-14 x 5.5-7 µ, with a slightly eccentric pore; growing alone, widely scattered, or in loose groups on sticks, litter, grass, wood chips, and muck.
Coprinellus plagioporus

75.Not completely as above.

76.Pileocystidia variable in shape, frequently with thick, yellowish to brownish walls; spores 10.5-15 x 5-7.5 µ, with an eccentric pore; growing scattered to gregariously or in clusters on soil rich with litter or woody debris, especially in disturbed-ground locations, or on lawns; in North America suspected from Idaho.
Coprinellus sclerocystidiosus

76.Not completely as above.

77.Usually found near woody debris (including wood chips) in disturbed-ground areas like paths and ditches; cheilocystidia exclusively subglobose; spores 6-7.5 µ wide; some or many pileocystidia with thick walls present.
Coprinellus callinus

77.Usually found in naked soil, muck, or mull; cheilocystidia often polymorphic, ranging from subglobose to lageniform; spores 6-8 µ wide; thick-walled pileocystidia rare or absent.
Coprinellus subimpatiens

78.Basidia 2-spored.
Coprinellus amphithallus

78.Basidia 4-spored.

79.Growing gregariously on fallen sticks near Mt. Hood and possibly not collected since the 1947 type collection; pileocystidia short (40-70 µ long), with tapering necks; spores very broadly elliptical, measuring 8.5-10.5 x 7-8.5 µ.
Coprinellus eurysporus

79.Habitat various; pileocystidia longer and spores less broadly elliptical than above.

80.Growing in dense clusters in grass; veil present as chained, subcylindric to subfunky elements (scalp-section the disc); pileocystidia lageniform, tapering, always with thin walls; spores 7.5-11.5 x 4.5-6 µ.

80.Growing alone, scattered, or in loose clusters in bare soil or grass; veil elements absent; pileocystidia tapering or not, sometimes with thick walls; spore measurements various.

81.Pileocystidia cylindric or tapering only slightly, with thick-walled pileocystidia usually present; spores 9-14 x 6-8 µ, with an eccentric pore.
Coprinellus subimpatiens

81.Pileocystidia tapering substantially, always with thin walls; spores 10.5-11.5 x 6-7.5 µ, with a central or slightly eccentric pore.
Coprinellus impatiens

82.Spores 13-19 x 7-9.5 µ, with an eccentric pore; cap yellowish when young; veil with subglobose elements mixed in with tubular and sausage-shaped elements.
Coprinellus flocculosus

82.Spores shorter than above, with or without an eccentric pore; young cap variously colored; veil elements mostly tubular to sausage-shaped, rarely subglobose.

83.Spores with a sheathlike outer covering.

83.Spores without a sheathlike outer covering.

84.Spores subglobose to broadly lemon-shaped, 7-9 x 6-7.5 µ; growing alone in a Washington lawn in 1968 and now preserved as "[o]nly a stipe, a part of stipe with collapsed pileus as a black mass at apex and a small part of a pileus" (Uljé), and possibly otherwise undocumented.
Coprinopsis brunneistragulata

84.Spores elliptical, 11-15 x 7.5-9 µ; growing in a Washington greenhouse in 1950 and possibly otherwise undocumented.
Coprinopsis tectispora

85.Spores subglobose to very broadly elliptical or broadly lemon-shaped.

85.Spores more narrowly elliptical.

86.Growing as a pest in commercial mushroom farm compost; spores globose to subglobose.
Coprinopsis undulata

86.Growing elsewhere (often near woody debris in woods, or in burned areas); spores subglobose to broadly lemon-shaped.
Coprinopsis lagopides

87.Veil present as a dense covering of white hairs over a mouse gray cap surface; spores 10.5-13.5 x 7-8.5 µ, with a central pore; common and widely distributed in North America.

87.Veil and cap not colored as above (gray-on-gray, tan-on-brown, or colorless-on-brown instead); rare species apparently known only from type collections in the Pacific Northwest (though one species has since been found in England).

88.Veil scant, even in buttons, visible as "minute reddish brown scales or filaments"; spores 14-15 x 7-8 µ, with an apical pore; growing near woody debris of conifers.
Coprinopsis sylvicola

88.Veil more prominent than above, either colorless or tan; spores 9-11.5 µ long, with an apical or eccentric pore; habitats various.

89.Cap with tan veil on a brown surface; spores with an eccentric pore.
Coprinopsis alutaceivelata

89.Cap with colorless to brownish veil on a whitish surface that becomes brownish; spores with an apical pore.
Coprinopsis pachyderma


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Kuo, M. (2008, February). Coprinoid mushrooms: The inky caps. Retrieved from the MushroomExpert.Com Web site: