Bull. Natn. Sci. Mus., Tokyo, Ser. B, 31(4), pp. 117–126, December 22, 2005

Three Coprinus Species Occurred on the Animal Dungs Collected at
Yatsugatake Range, Central Honshu, Japan
Toshimitsu Fukiharu1, Kohtaro Takarada2, Tsuyoshi Hosoya3 and Noriko Kinjo4
1

Natural History Museum & Institute, Chiba, Aoba-cho 955–2, Chiba, 260–8682 Japan
E-mail: fukiharu@chiba-muse.or.jp
2
Faculty of Science, Toho University, Miyama 2–2–1, Funabashi, Chiba, 274–8510 Japan3
Department of Botany, the National Science Museum, Amakubo 4–1–1, Tsukuba, 305–0005 Japan
4
College of Liberal Arts and Sciences, Tokyo Medical and Dental University, 2–8–30,
Kohnodai, Ichikawa-shi, Chiba, 272–0827 Japan

Abstract Three Coprinus species i.e., Coprinus candidolanatus, C. stercoreus and C.
ephemeroides occurred on animal dungs collected in Yatsugatake were documented. Production ofbasidiomata of Coprinus stercoreus and C. candidolantus started in 8 and 10 days, respectively,
followed by C. ephemeroides, and lasted for about 40 days. At the maximum, 30 basidiomata were
produced from ten dung particles. Descriptions and illustrations together with proper binomials to
the two species and comments on the third species are given.
Key words : Coprinus candidolanatus, Coprinus ephemeroides, Coprinus stercoreus, dungfungi, mycobiota.

Introduction
Dung of herbivorous animal is a good substrate for various fungi, the life cycles of whichlargely depend on dung. These fungi are called“dung fungi” or “coprophilous fungi” (Bell,1983; Furuya, 1990; Richardson, 2001; Webster,1970). Dung fungi are composed of myxomycetes, zygomycetes, ascomycetes, basidiomycetes, and anamorphic fungi, and they occur
in succession on dung (Richardson, 2002). Although dung fungi have been investigated formore than 100 years in Europe (Cailleux, 1971;
Massee and Salmon, 1901, 1902; Moreau, 1953)
and North America (Cain, 1934; Griffiths, 1901),
still new members, including new taxa are being
added (Uljé and Keizer, 2003; Uljé and Noordeloos, 2000; Uljé et al., 2000).
In Japan, coprophilous zygomycetes (Indoh,
1962, 1965), yeasts (Soneda, 1959, 1962), discomycetes (Minoura and Yamada, 1976; Minoura
et al., 1978; Otani, 1973; Otani and Kanazawa,
1970a, 1970b), pyrenomycetes (Furuya and Udagawa, 1972a, 1972b, 1973; Udagawa and Furuya,
1974, 1977) and anamorphic fungi (Tubaki,1954) have been studied. In spite of rich fungal
diversity in Japan, research on dung fungi has
relatively shorter history than that in Europe, andstill studies of mycobiota are required. In particular, studies on coprophilous basidiomycetes are
scarce.
Coprinus Pers. is the most dominant genus
among the coprophilous basidiomycetes. A large
number of Coprinus species are known to occur
on low C/N ratio materials, such as ammonium
treated soils (Sagara, 1975, 1992), compost, anddung (van Waveren, 1968), dung being one of thebest substrates. Because the basidiomata of Coprinus degrades very quickly in the field, it isusually difficult to obtain specimens in good condition. On the other hand, occurrence of Coprinus on dung can be monitored under controlled
condition by continuous observation, and specimens in good conditions can be obtained andpreserved. This gives an advantage in taxonomical studies.
About 40 species of Coprinus are hitherto
known in Japan, which include 7 coprophilous
species (Hongo, 1987). However, literature sur-

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vey suggests that 16 species of them may be included as coprophilous members in Coprinus(Breitenbach and Kränzlin, 1995; Moser, 1983).
During the taxonomic research of flora in areawith remarkable biodiversity in Japan, Yatsugatake area, Nagano prefecture was surveyed forthe occurrence of coprophilous basidiomycetes.
In the present paper, the authors provided proper
binomials, detailed descriptions and illustrations
for two Coprinus species in Japan, and comments on the third species.
Materials and Methods
Dung of sika deer (Cervus nippon) and Japanese serow (Capricornis crispus) were collected in
two sites at the foot of Yatsugatake range, Chino,
Nagano Prefecture on Aug. 12 and Oct. 13 in
2004. Six specimens were collected for eachspecies. The collected specimens were brought tothe laboratory in paper bags, divided to two portions, and each was incubated on moistened mossin tall Petri-dishes at 15°C or 22°C under a 12hours light/12 hours dark light regime. The spec-

imens were examined every 3–4 days for the occurrence of fungi under dissecting microscope(Fig. 1). Basidiomata were counted for the frequency data, collected for identification and used
for isolation. To isolate culture, a small portionof the young basidioma was taken by a sterilized
needle, and transferred onto a medium preparedin a test tube. Cultures were grown at 22°C undera 12 hours light/12 hours dark light regime on
MYC medium (malt extract 10 g, yeast extract
2 g, casamino acid 2 g, agar 12 g, D. W. 1L; Difco
Laboratories) for about 2–3 weeks, and matured
basidiomata were collected for identification andfreeze-dried for herbarial specimens. Morphological observations of the anatomy and measurements were made from freeze-dried specimens
mounted in 25% aqueous ammonia. Descriptionsof macro- and microscopic characters were prepared based on basidiomata both cultivated on
artificial medium and on the dung specimens.
Specimens examined are deposited in the National Science Museum, Tokyo (TNS) and Natural
History Museum and Institute, Chiba (CBM).

Fig. 1. Moist chamber for dung incubation. A. Dungs in a moist chamber. Moistened moss were seated under
the dung to keep the chamber in moist condition. B. Dung fungi were examined under dissecting microscope.

Coprinus spp. on dung

Results and Discussion
Before the appearance of the basidiomycetes,
zygomycetes, anamorphic fungi, and ascomycetes occurred in this order on dung specimens. The successive occurrence of fungi lasted
for up to 10 weeks and 14 weeks at 22 and 15°C,
respectively. All the basidiomycetes occurred belonged to the genus Coprinus. The occurrence
and the occurring period differed by the species
and incubation condition. Seven taxa of Coprinus
were recognized, the three of which were identified: Coprinus candidolanatus, C. stercoreus,
and C. ephemeroides. The frequency of occurrence and incubation condition are listed in Table
1. Production of basidiomata of Coprinus stercoreus and C. candidolantus started in 8 and 10
days, respectively, and lasted for about 40 days.
At the maximum, 30 basidiomata were produced
from ten dung particles (Fig. 6). Coprinus candidolanatus and C. stercoreus were observed only
under 22°C while C. ephemeroides occurred atboth temperatures. The identification of the remaining 4 taxa are in process. Two to three
species of Coprinus species occurred from eachspecimen, but overlapping species were few.
Descriptions
Because Coprinus candidolanatus and C. stercoreus are identified and provided proper binomials for the first time in Japan, detailed descriptions are given as follows.
Coprinus candidolanatus Doveri & Uljé in
Uljé, Doveri & Noordeloos, Persoonia 17:465. 2000.
Figs. 2, 3
Pileus 1–2 mm broad, 2–3 mm high when closed,
ellipsoid to ovoid when young, becoming convex
to plane, 3–6 mm broad when expanded, turning
applanate at maturity or even revolute at deliquescence; ground color at first white, soon becoming pale grayish ochre, surface densely covered with a fibrous-wooly white universal veilwhen young (Fig. 2B), sometimes with cortinate
veil in very young stage (Fig. 2C), later almost
glabrous or veil remaining only at the center.
1.

119

Flesh very thin, fragile, white, taste mild, odorless. Lamellae free, dispersed (number of lamellae reaching stipe25–30), narrow (0.4–0.7 mm
breadth), at first white, then grayish, finally
blackish, deliquescent. Stipe up to 2–5 cm
0.5 mm (Fig. 2A), cylindrical, equal, sometimes
with clavate base, not rooting, fragile; surface
white, at first with white fibrillose or squarrose
scales, soon becoming smooth. Basidiospores
black in mass, dark red-brown under the microscope, ovoid to ellipsoid, smooth, 8.5–10.0(9.170.51: meanSD, n40) m m long, 4.7–5.9(5.430.27, n20) m m broad in face view,
4.7–5.9 (5.410.26, n20) m m in side view, acentral germ pore 1.2–1.5 m m wide (Figs. 2H,
3D). Cheilocystidia 35–70 m m long, 20–30 m m
broad, ellipsoid to obovoid, numerous, thinwalled, hyaline (Fig. 2F, 3B). Pleurocystidia
45–65 m m long, 24–27 m m broad, ellipsoid, thinwalled, hyaline (Figs. 2G, 3C). Veil on the pileal
surface composed of two type of thin-walled hyaline hyphal element: chain of sausage-shaped hyphae without branch (20–20010–35 m m) andcluster of diverticulate hyphae with small sidebranches and branchlets (Figs. 2D, 2E, 3A).
Clamp-connection not recognized.
Habitat. On dungs of Japanese serow(Capricornis crispus), also on the dung of cow
and pig (Aoki, 1981), on dungs of deer and sheepin the field (Uljé et al., 2000).
Known distribution. Italy and Netherland(Uljé et al., 2000), Kyoto and Saitama Prefecture(Aoki, 1981), Tokyo (Aoki, 1993).
Specimens
examined. CBM-FB-34890
TNS-F-11630,
CBM-FB-34950,
CBM-FB35061, CBM-FB-35062, CBM-FB-35768 (on thedung); CBM-FB-35441TNS-F-11631 (basidiomata produced in culture). All specimens werefrom the dung of Japanese serow collected in
Yatsugatake, Chino, Nagano Prefecture, alt. 2000
m in Aug. 12, 2004.
Japanesename. Shirage-ushiguso-hitoyotake (Aoki, 1981, 1993).
Notes. The isolate produced mature basidiomata on MYC medium. The fungal specimen obtained in the present study differed a little from

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Fig. 2. Coprinus candidolanatus. A. Basidiomata on the dung specimens in moist chamber after 18 days of incubation. B. Basidiomata cultivated on MYC medium in 10 days, FB-35441 (CBM). C. Basidiomata on thedung in moist chamber of 9 days incubation. D, E. Veils on the pileipellis. The element of the veil was composed of two kind of hyphae: sausage-shaped elements (indicated by white arrows) and diverticulate elements(indicated by black arrows), FB-35441 (CBM). F. Cheilocystidia, FB-34890 (CBM). G. Pleurocystidia, FB34890 (CBM). H. Basidiospores, FB-34950 (CBM). Scales. A2 cm, B5 mm, C2 mm, D,E50 m m, F,
G20 m m, H10 m m.

the holotype in the size of basidiomata (holotype,pileus 3–62–4 mm when closed) and in the proportion of the basidiospore (length/bredthca. 1.5). However, conspicuous character of veilon the pileal surface, i.e. veil composed of two
type of element, chain of sausage-shaped andcluster of diverticulate element with side-branches and branchlets are the unique characters to

distinguish this species from others in this genus(Uljé et al., 2000). Aoki (1981, 1993) documented this unique fungus as Coprinus sp. in his private communications in Japanese, giving a newJapanese name to it. Because the authors identified the present fungus as C. candidolanatus andrecognized that it was identical to Aoki’s Coprinus sp. (Aoki 1981, 1993), we document the fun-

Coprinus spp. on dung

121

Fig. 3. Coprinus candidolanatus. A. Veils on the pileipellis, FB-35441, 34890 (CBM). B. Cheilocystidia, FB34890 (CBM). C. Pleurocystidia, FB-34890 (CBM). D. Basidiospores, FB-34950 (CBM). E. Basidiomata on
the dung in moist chamber of 9 days incubation. Scales. A–C20 m m, D10 m m, E2 mm.

gus here, adopting Aoki’s Japanese name, and his
observations were added to the previously known
distribution in Japan. In his private communications, Aoki (1981, 1993) also described two otherCoprinus spp. with two or three hyphal elemetnt
in the pileal veil, but did not assign any name tothem. These species differ from C. candidolanatus in the spore shape and veil element type.
Coprinus stercoreus Fr. Epicrisis systematis
mycologici (Uppsala) p. 251. 1838.
Figs. 4, 5
Coprinus stercorarius (Bull.) Fr. Epicr. syst.
mycol. p. 251. 1838.
Coprinus velox Godey apud Gillet, Cham. Fr.
Hym. p. 614. 1878.
Coprinus evanidus Godey apud Gillet, Cham. Fr.

2.

Hym. p. 614. 1878.
Coprinus velox var. stenosporus Svrček, Česká
Mycol. 10: 176. 1956.
Coprinus stercoreus Fr. sensu Watling, Notes R.
bot. Gdn Edinb. 28: 48. 1967.
Coprinus velox Godey sensu Kits van Waveren,
Persoonia 5: 154. 1968.
Coprinopsis stercorea (Fr.) Redhead, Vilgalys &Moncalvo, in Redhead, Vilgalys, Moncalvo,
Johnson & Hopple. Taxon 50: 231. 2001.
Pileus 1.5–2 mm broad, 2–3 mm high whenclosed, when young ellipsoid to ovoid, later campanulate to plane, 3–4 mm broad when expanded,turning applanate at maturity or even revolute atdeliquescence, ground color at first white, soon
becoming pale greyish ochre, surface when
young densely covered with a wooly white uni-

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Fig. 4. Coprinus stercoreus. A, B. Basidiomata cultivated on MYC medium in 14 days, FB-35766 (CBM). C.
Basidiomata cultivated on MYC medium in 17 days, FB-35415 (CBM). D, E, F. Veils on the pileipellis, theelement of the veil was composed of two kind of hyphae, sphaerocyst elements and diverticulate elements,
FB-35414 (CBM). G. Cheilocystidia, FB-35414 (CBM). H. Pleurocystidia, FB-35414 (CBM). I. Basidiospores, FB-35767 (CBM). Scales. A1 cm, B5 mm, C5 mm, D, E50 m m, F100 m m, G, H
20 m m, I10 m m.

versal veil (Fig. 4A, B), consisting of a whitish
granules mixed with a wooly hairy mass of fibers
particularly near the margin of the cap, later theveil remaining only in the center. Flesh very thin,fragile, white, taste mild, odorless. Lamellae
free, dispersed (number of lamellae reachingstipe20–30), narrow (0.3–0.6 mm breadth), at
first white, then grayish, finally blackish, deliquescent. Stipe up to 2–5 cm0.5 mm, cylindrical, equal, sometimes the base clavate, not rooting, fragile, surface white, at first with white fibrillose scales, soon becoming smooth. Ba-

sidiospores black in mass, dark red-brown underthe microscope, ellipsoid, often phaseoliform,
smooth, 6.3–7.5 (6.720.28: meanSD,n40) m m long, 2.8–3.8 (3.330.22, n20) m m
broad in face view, 3.1–3.4 (3.230.15,n20) m m in side view, a central germ pore 1.1–
1.3 m m wide, apiculus excentric on the adaxial
face (Figs. 4I, 5D); perispore not observed.
Cheilocystidia 20–35 m m long, 13–23 m m broad,
ellipsoid to obovoid, numerous, thin-walled, hyaline (Fig. 4G, 5B), Pleurocystidia 17–23 m mlong, 12–15 m m broad, subellipsoid to obovoid,Coprinus spp. on dung

123

Fig. 5. Coprinus stercoreus. A. Veils on the pileipellis, the element of the veil was composed of two kind of
hyphae, sphaerocyst elements and diverticulate elements, FB-35414 (CBM). B. Cheilocystidia, FB-35414(CBM). C. Pleurocystidia, FB-35414 (CBM). D. Basidiospores, FB-35767 (CBM). Scales. A–C10 m m, D
5 m m.

thin-walled, hyaline (Figs. 4H, 5C). Universal
veil on the pileal surface composed of two typeof element; verrucose globose to subglobose
cells (diameter; 20–40 m m, young specimen) andcluster of diverticulate hyphae with side-branches and branchlets (diameter; 3–5 m m, Figs. 4D–F,
5A). Clamp-connection not recognized.
Habitat. On dung of Japanese serow (Capricornis crispus) or dung of rabbit (Lepus brachyurus, Aoki, 1980), on dung of cow, horse, sheep,
rabbit, deer (Kits van Waveren, 1968).
Known distribution. Europe, North America, North Africa (Breitenbach and Kränzlin,
1995), Aichi (Aoki, 1981).
Specimens
examined. CBM-FB-35079,
CBM-FB-35080, CBM-FB-35767 (on dung ofJapanese serow collected in Yatsugatake range,
2,000 m elevation on August 12, 2004); CBMFB-35414,
CBM-FB-35415TNS-F-11632,

CBM-FB-35766TNS-F-11633(basidiomata
produced in culture).
Japanese name. Tofun-hitoyotake (Aoki,
1980).
Notes. Coprinus stercoreus belongs to the section Vestiti (Lange) Khner & Romagnesi (Moser,
1983; Singer, 1986) because of its sphaerocyst
element in the pileal veil. In this section two
species have been described: Coprinus tuberosus
Qul. and C. stercoreus both of which have no or
few perispore on the surface of the basidiospores,
with two types of pileal veil element (sphaerocysts and filamentous hyphae with small sidebranches and branchlets), spore length under
10 m m (Breitenbach and Kränzlin, 1995; Moser,
1983; Orton and Watling, 1979; van Waveren,1968). Coprinus stercoreus is distinguished fromC. tuberosus in the character of basidiospore(length under 9.5 m m, often phaseoliform) and124

T. Fukiharu et al.

having no sclerotium under the stipe (Breitenbach and Kränzlin, 1995; van Waveren, 1968).
The idea to transfer the present fungus to Coprinopsis (Redhead, 2001) is currently not widely
Table 1.

accepted, and we followed the conventional taxonomy. Hongo (1987) documented C. stercorarius from Japan, but we determine that his specimen was conspecific with C. tuberosus based on

Occurrence of the three Coprinus species documented in the present study from the dung specimens.
Origin of the dung

Incubatedtemperature(°C)

No. ofspecimens

15

6

0

0

0

22

6

0

0

1

15

6

0

0

1

22

6

1

1

0

Coprinus
Coprinus
Coprinuscandidolanatus stercoreus ephemeroides

sika deer (Cervus nippon)

Japanese serow (Capricornis crispus)

Fig. 6. Number of basidiomata of Coprinus spp. on the dung specimens. A. Coprinus candidolanatus () andC. stercoreus () on the same dung of Capricornis crispus, incubated at 22°C for 9 weeks. B. Coprinus
ephemeroides () on the dung of Capricornis crispus, incubated at 15°C for 14 weeks. C. Coprinus
ephemeroides () on the dung of Cervus nippon, incubated at 22°C for 14 weeks.

Coprinus spp. on dung

his description. It seems that Aoki (1980) was thefirst to recognize the present fungus in Japan, but
he identified it as C. velox Godey and published
it in his private communication in Japanese withJapanese name. Because the authors recognizedthat the present fungus was identical to Aoki’s
description, we describe and illustrate it here
under the current name, adopting Aoki’s Japanese name, respecting his work. The present fungus is fairly common as a coprophilous fungus in
the world (van Waveren, 1968).
3.

Coprinus ephemeroides (Bull.) Fr.
Habitat. On dungs of various kinds (Orton
and Watling, 1979); on dungs of cow and horse,
Tatami mat disposed in the field (Hongo, 1987),
on the dung of Cervus nippon and Capricornis
crispus.
Known distribution. North America, Europe (Moser, 1983; Orton and Watling, 1979) ;Otu (Hongo, 1972), Tokorozawa (Aoki, 1976,
1995).
Specimens examined. On dung of sika deer
(Cervus nippon); CBM-FB-35438, CBM-FB35440, CBM-FB-35443TNS-F-11634. On
dung of Japanese serow (Capricornis crispus);
CBM-FB-35439TNS-F-11635,
CBM-FB35442, CBM-FB-35444, CBM-FB-35446TNSF-11636. All dung specimens were collected in
Yatsugatake, Chino, Nagano Prefecture in Oct.13
2004.
Japanesename. Tsuba-hina-hitoyotake(Hongo, 1972)
Notes. Morphological characteristics agreed
with those given by Hongo (1972, 1987). In thepresent study, C. ephemeroides occurred on dungof Cervus nippon (incubated at 22°C) and Capricornis crispus (incubated at 15°C). Isolationfrom the fruiting body was unsuccessful.
Coprinus ephemeroides belongs to the section
Vestiti (Lange) Khner & Romagnesi (Moser,
1983; Singer, 1986) because of its sphaerocyst
element in the pileal veil. Coprinus
ephemeroides is relatively well-documented partly because it is easily distinguished from otherspecies and can be identified by combination of125

obvious characters such as ring on the stipe,
without perispore on the surface of the basidiospores which are pentagonal in face view.
Acknowledgements
The author expresses their gratitude to Dr.
Keiji Ochiai, (Natural History Museum & Institute, Chiba) for identification of the dung. They
thank Ms. Haruko Suga (Chiba mycologicalclub) for her assistance in collecting the specimens. Mr. Ichiro Gomi, Suwa City Museum of
Art is greatly appreciated for his excellent guiding in the field. The authors wishes to express
their gratitude to Dr. Y. Hiratsuka for reviewing
the manuscript.
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