Fig. 1. Cheilymenia stercorea, apothecium.
Fig. 2. C. stercorea, stellate and rooted hairs.
Fig. 3. Lasiobolus ciliatus, apothecium.
Fig. 4. Iodophanus carneus, apothecium and spore.
| Transcriber's note: | Corrigendum applied at the wish of the principle author: in Key 3 the pointers to couplets 56 and 66 were the wrong way round and have been corrected in this edition. |
KEYS TO FUNGI
ON DUNG
by
M. J. RICHARDSON
165 Braid Road,
EDINBURGH EH10 6JE
and
ROY WATLING
Royal Botanic Garden,
EDINBURGH EH3 5LR
Published by the British Mycological Society
PO Box 30, Stourbridge
West Midlands DY9 9PZ
© British Mycological Society 1997
Printed in Scotland by BPC-AUP Aberdeen Ltd
ISBN 0 9527704 2 3
The first edition of these keys was published in the Bulletin of the British Mycological Society 2, 18-43 (1968) and 3, 86-88, 121-124 (1969) in an attempt to bring together in one place information for the identification of coprophilous fungi which would be useful to teachers and others interested in these fungi. They were issued as a separate publication in 1972, and with corrections in 1974. They were reprinted in 1982 with additions. This latest edition is an update of all the earlier ones, with current nomenclature and recent references, and the inclusion of some additional species.
M.J.R.
R.W.
December 1996
Coprophilous fungi are highly satisfactory for demonstrating the diversity and morphology of a group of related organisms within an ecological system. Representative genera of most major groups of fungi can usually be guaranteed to appear on dung after a period of incubation. There is no shortage of dung in our fields and woods, and this material will always produce characteristic fungi at whatever time of year it is collected.
Dung is best incubated in a light place, for example on a table in a warm room, on layers of moist filter paper or other absorbent material. For rabbit pellets, and samples of similar size, Petri dishes are ideal; for horse 'apples', and larger types of dung, large covered dishes such as glass casseroles, plastic sandwich boxes or yoghurt pots are needed. The top third cut from a plastic lemonade or mineral water bottle fits neatly in a Petri dish, and replacing the screw cap with a cotton wool plug allows aeration and gives adequate height for developing basidiomycetes. Samples should not be kept in airtight containers for any length of time after collection, as in such conditions insects and nematodes tend to break down the dung, and anaerobic conditions which do not favour the fungi rapidly develop. If they cannot be set to incubate soon after collection they can be gently air dried, as most dung fungi will remain alive after such treatment and grow out when the sample is eventually moistened. The absorbent material should be kept moist. Although free water will not allow the best development of ascomycetes, the succession of basidiomycetes appears to vary with the wetness of the dung. Earthworms and insect larvae should be excluded from the samples as far as possible, for they break up the dung too much; activity of the latter can be reduced by spraying lightly with a household insecticide. If space is limited and cultures are kept nearby, it is very important to prevent mite infestation. Containers can be isolated by placing on glass plates lightly smeared with Vaseline, to which an acaricide (e.g. methyl benzoate) can be added.
Fungi are best sought with a stereoscopic binocular microscope, when their full beauty will be seen, but a hand lens or simple magnifier, although less convenient, is sufficient for all but the smallest forms. The larger ascomycetes and most of the basidiomycetes are readily seen with the unaided eye, but the binocular microscope is still very useful for observing the gross features of the veil of the basidiomycetes. Perithecia, apothecia and similar structures can be removed with fine needles or forceps quite cleanly for mounting, initially in water, on slides. Subsequent irrigation with iodine solution will allow any reaction of ascus wall, tip or pore to be observed, and mounting in diluted Indian ink can enhance the visibility of appendages, caudae and sheaths which occur on some spores. Spore discharge in the ascomycetes often occurs from mature asci when material is mounted in water, so mature spores can immediately be seen. Many of the coprophilous toadstools (agarics), because of their small size and/or rapidly deliquescent nature, often do not give spore prints in the normal way, but mature spores can usually be found on the stipe or in natural spore prints formed on the absorbent material on which the dung is supported. For accurate identification the ability to measure the size of spores and other structures will be necessary. Basic microscopical technique and mycological knowledge is assumed. Common species are well described and illustrated in popular books, and references are given to specialist works to allow descriptions of less common species to be found. It will be necessary to refer to these for critical taxa. Although this edition contains about one half more species than the 1982 edition, there are still many species to be described and new records and observations to be made, especially in the Ascomycotina.
Four keys are presented. Keys 1 and 2 (MJR) are to the coprophilous ascomycetes, a very diverse group which, although not covering all the possible types of reproductive structure found in the class, contains many of the important types. The information for the identification of these fungi is dispersed throughout the literature, and many new species are still being discovered and described. Some appear to be world-wide in their distribution, others more restricted, with a prevalence of reports from either arctic, temperate or tropical regions. These keys are not exhaustive, since there are far too many species to make it practical to include them all. They do, however, include most genera, and the commoner or well known species of temperate regions. Specific (and even generic) limits in some cases (e.g. Coprotus / Ascophanus / Ryparobius / Thelebolus) are still the subject of debate and the choice of names to use in the key for a few taxa has been a compromise. Key 2 includes the original 'plectomycete' key (RW), which contains fungi which may not be strictly coprophilous in the normal sense, but fungi which occur on hair, horn, bone and cadavers, and may thus be found on carnivore dung or pellets of owls and other birds of prey.
Key 3 (RW, p. 52) is to the basidiomycetes of dung and associated debris. The part of the key dealing with the agarics attempts to be as complete as possible. Since the toadstools have always been thought of as the best known of the coprophilous fungi, attention to their taxonomy has often been careless. In this key the opportunity has been taken to adopt a rather narrow species concept, and to provide in certain places indications of where distinct taxa, even autonomous species, may be found after further laboratory work. Many of these types have been cultured and appear to differ vegetatively in ways which support observations of gross morphology. Coprophilous agarics are popular material for genetic studies and additional information on veil structure, spore number etc. of individual species is given, even when these are not 'key characters'.
Key 4 (MJR, p. 63) is to the Zygomycota (phycomycetes) which are characteristic of dung and amongst the first to appear when freshly dropped dung is incubated. They soon disappear, however, but their fruiting can be prolonged by plating small portions of dung on a nutrient medium (e.g. potato carrot or potato dextrose agar) to which has been added a small amount of antibiotic to reduce bacterial growth. This method is especially suitable for the parasitic and predacious fungi. A cultural approach is essential for the identification of many of these fungi and the above media, and oatmeal agar, are suitable for culture as well as isolation. For this reason the study of this group of fungi is less easy than that of the ascomycetes and basidiomycetes but, because the asexual stages are characteristic, we have attempted to key out the commoner genera which might be found, with notes on common species. The asexual spores are sporangiospores formed in sporangia; some sporangia produce a single spore within a closely fitting sporangium, and have in the past been erroneously described as conidia. A great range of sporangial structure occurs within the orders concerned. The classical structure is the massive (up to 250µm diam.) multispored sporangium with an internal columella which remains after the spores have been dispersed (e.g. Mucor); those of Mortierella are similar, but smaller and without a columella. Other sporangia are much reduced and may be only 10-20µm diam., and contain only a small number of spores (Thamnidium) or one spore (Chaetocladium); these small globose structures are termed sporangioles. Spores may also form in chains; the chains are in terminal groups and are formed by the differentiation of the contents of cylindrical sporangia which are considered to be part-sporangia (merosporangia). When the sporangial wall has disappeared the spore chains may remain discrete and intact, or they may collapse into a wet droplet of spores (Syncephalastrum, some Piptocephalis). Members of the Kickxellaceae (e.g. Coemansia, Kickxella) have single spored merosporangia produced in serried ranks on boat-shaped or swollen structures (sporoclades). The sexual spores (zygospores) are rarely seen without culturing; oatmeal agar is one which favours their production. The key includes one member of the Entomophthorales, which also produces single-spored sporangia. Other members of this order may be found parasitising the various animals which live in dung; many other predacious fungi may also be seen, e.g. parasites of amoebae (Acaulopage). The key is of necessity far from complete, and omits members of the Dimargaritales, which have been found frequently on dung of small mammals in America.
Mitosporic fungi ('Fungi Imperfecti') and myxomycetes have been excluded, since they would expand the range of these keys beyond what was initially intended, although numerous species of both groups occur on dung when incubated in a damp chamber. For mitosporic fungi see Seifert, Kendrick & Murase (1983) and Ellis & Ellis (1988); for myxomycetes see Eliasson & Lundqvist (1979). As practical keys, rather than a taxonomic treatment, taxonomic authorities have not been cited. For ascomycetes, Cannon, Hawksworth & Sherwood-Pike (1985) have been followed, unless there is a more recent treatment of a group. For the basidiomycetes the 'New Checklist of British Agarics and Boleti' (Dennis, Orton & Hora, 1960, Supplement to the Transactions of the British Mycological Society 43) has been followed, and The British Fungus Flora (Orton & Watling, 1979 and Watling, 1982).
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BASIDIOMYCETE REFERENCES
Moser, M. (1978), in Gams, H. (ed.). Kleine Kryptogamenflora von Mitteleuropa. Fischer Verlag.
Moser, M. (1983). Keys to Agarics and Boleti (English translation by S. Plant). Roger Phillips, London.
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PHYCOMYCETE REFERENCES
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GENERAL REFERENCES
Bell, A. (1983). Dung Fungi: an illustrated guide to coprophilous fungi in New Zealand. Victoria University Press, Wellington.
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Cacialli, G., Caroti, V. & Doveri, F. (1995). Funghi fimicoli e rari o interssanti del litorale Toscano. Schede di Micologia vol. 1. Fondazione Centro Studi Micologici Dell' A. M. B., Vicenza, Italy.
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Ellis, M. B. & Ellis, J. P. (1988). Microfungi on Miscellaneous Substrates. Croom Helm, London & Sydney.
Gilman, J. C. (1957). A Manual of Soil Fungi. Iowa State College Press.
Eliasson, U. & Lundqvist, N. (1979). Fimicolous Myxomycetes. Botaniska Notiser 132, 551-568. (A list of 34 spp., with some descriptions and illustrations).
Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler, D. N. (1995). Ainsworth & Bisby's Dictionary of the Fungi. 8th edn. CAB International, Wallingford.
Holden, M. (ed) (1982). Guide to the literature for the identification of British fungi, 4th Edition. Bulletin of the British Mycological Society 16, 36-55; 92-112.
Massee, G., & Salmon, E. S. (1901). Researches on coprophilous fungi. Annals of Botany, London 15, 313-357.
Seifert, K. A., Kendrick, W. B. & Murase, G. (1983). A key to hyphomycetes on dung. University of Waterloo Biology Series No. 27.
Webster, J. (1970). Coprophilous Fungi. Transactions of the British Mycological Society 54, 161-180.
| 1 | Ascoma either globose to flask shaped, usually with an easily observable pore or neck (perithecium or pseudothecium, figs 16, 18, 19, 22, 27, 30, 32, 34-37), or discoid (apothecium, figs 1, 3, 4, 7, 11-14). Spores usually 8 in each ascus (less frequently 4, 16, 32, 64, 128 etc.). Asci ellipsoid to cylindrical, borne in a distinct hymenium, thus appearing in fascicles or distinct groups when the fruit body is squashed. | 2 |
| - | Ascoma globose to subglobose, lacking a definite pore or neck (cleistothecium or gymnothecium, figs 38, 39, 46). Asci globose to subglobose, 8-spored, not in a distinct hymenium, appearing quite free when the fruit body is squashed. | |
| Key 2, 148 (p. 45) | ||
| 2(1) | Ascoma a perithecium or pseudothecium, usually dark in some part, not opening to a disc but remaining globose or flask shaped. Asci unitunicate, not operculate but often with an apical pore, which may stain blue in iodine, or bitunicate. | |
| Key 2, 1 (p. 24) | ||
| - | Ascoma an apothecium, white or lightly coloured, soft fleshed, opening out to a disc or cushion shape when mature. Asci unitunicate. | 3 |
| 3(2) | Asci opening by an operculum (fig. 8), a bilabiate vertical split down to a subapical ring of thickening (fig. 15), or apparently just bursting. | 4 |
| - | Asci inoperculate, with an apical pore. | 96 |
| 4(3) | Spores 8 (occasionally 4) in an ascus, colourless, purple or brown. | 5 |
| - | Spores more than 8 in an ascus, colourless. | 77 |
| 5(4) | Spores remaining colourless. | 6 |
| - | Spores purple or brown at maturity. | 39 |
| 6(5) | Apothecia with obvious hairs. | 7 |
| - | Apothecia without obvious hairs (microscopic hairs up to 50µm long may be present). | 14 |
| 7(6) | Hairs brown. Apothecia orange, red orange or yellow orange | |
| (Cheilymenia, fig. 1) 8 | ||
| - | Hairs colourless. Apothecia colourless or pinkish. | |
| (Lasiobolus, fig. 3) 12 | ||
| 8(7) | Apothecia with stellate hairs. Spores 14-20 × 8-11µm. | |
| Cheilymenia stercorea (figs 1, 2) | ||
| - | Apothecia without stellate hairs. | 9 |
| 9(8) | Spores 14.5-18 × 8-9.5µm. Asci 10-13µm diam. Apothecia 2mm diam. or more. | |
| Cheilymenia coprinaria | ||
| - | Spores larger, 17 × 10µm or more. | 10 |
| 10(9) | Apothecia reddish orange, up to 1mm diam., marginal hairs rooting, wall 2-4µm thick. Spores 21-26 × 10-13.8µm. | |
| Cheilymenia fimicola | ||
| - | Apothecia pale orange yellow, marginal hairs superficial, wall up to 2µm thick. | 11 |
| 11(10) | Asci up to 22µm diam. Spores 17-27 × 10-14.5µm. | |
| Cheilymenia pulcherrima | ||
| - | Asci wider, 25µm diam. or more. Spores 23-26.5 × 13-16.5µm. | |
| Cheilymenia raripila | ||
Fig. 1. Cheilymenia stercorea, apothecium.
Fig. 2. C. stercorea, stellate and rooted hairs.
Fig. 3. Lasiobolus ciliatus, apothecium.
Fig. 4. Iodophanus carneus, apothecium and spore.
| 12(7) | Hairs 600µm or longer. Spores 19-23 × 7-10µm. | |||
| Lasiobolus macrotrichus | ||||
| - | Hairs shorter, up to 600µm. | 13 | ||
| 13(12) | Asci clavate, 20µm diam. or wider. Spores 19-22 × 10.5-13.5µm. | |||
| Lasiobolus cuniculi | ||||
| - | Asci cylindrical, up to 20µm diam. Spores 18-22.5 × 9.5-11.5µm. | |||
| Lasiobolus ciliatus (fig. 3) | ||||
| 14(6) | Asci blue in iodine solution. | 15 | ||
| - | Asci not blue in iodine. | 24 | ||
| 15(14) | Spores large, 30-42 × 15-18µm, warted, ellipsoid with acute apices. | |||
| Thecotheus cinereus | ||||
| - | Spores smaller, smooth or only finely ornamented | 16 | ||
| 16(15) |
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| - | Apothecia pale, up to 4mm diam. Asci protruding from hymenium when ripe. | 17 | ||
| 17(16) | Apothecia white to pink, up to 2mm diam. Spores finely verruculose, 18-25 × 8-14µm. | |||
| Iodophanus carneus (fig. 4) | ||||
| - | Apothecia pale, variously coloured when fresh, but drying darker. Spores smooth. | |||
| (Thecotheus) 18 | ||||
| 18(17) | Spores apiculate at each end, smooth. | 19 | ||
| - |
|
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| 19(18) | Spores with a collar at the base of the apiculus. | 20 | ||
| - | Spores without a collar at the base of the apiculus, 16-21 × 8-12µm. | |||
| Thecotheus apiculatus | ||||
| 20(19) | Apothecia white. Spores 20-22 × 10-12µm, apiculus 4-6µm diam. | |||
| Thecotheus perplexans | ||||
| - | Apothecia yellowish. Spores 12-15 × 7.5-9µm, apiculus 2.5-3.5µm diam. | |||
| Thecotheus africanus | ||||
| 21(16) | Spores smooth, without guttules. | 22 | ||
| - | Spores verruculose or spinulose, 15-18 × 8-9µm, with 1 guttule. Paraphyses with clavate apices, with brown contents. Apothecia asymmetrical, extended on one side. | |||
| Peziza pleurota | ||||
| 22(21) | Spores 19-24 × 10.5-14µm. Apothecia yellowish brown, up to 10cm diam. | |||
| Peziza vesiculosa | ||||
| - | Spores up to 10µm wide. | 23 | ||
| 23(22) | Apothecia ca 1cm diam., umber with a paler margin. Spores 15-22 × 9-10µm. | |||
| Peziza bovina | ||||
| - | Apothecia up to 2cm diam., pale brown. Spores 13-16 × 7-9µm. | |||
| Peziza fimeti | ||||
| 24(14) | Apothecia robust, up to 4mm diam., orange or with brownish or purple tints. | 25 | ||
| - | Apothecia smaller, rarely more than 1mm, pale, yellowish green, orange, grey or chestnut. | 32 | ||
| 25(24) | Apothecia orange or red. | 26 | ||
| - |
|
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| 26(25) | Apothecia crowded, 1-3mm diam., orange, with a granular surface. Asci up to 190 × 15µm. Spores 15-18.5 × 7-9.5µm. Paraphyses strongly clavate to apex up to 14µm diam, filled with orange granules. | |||
| Coprobia granulata | ||||
| - | Apothecia discrete, 1-2mm diam., orange or red. Asci 240 × 10-12µm. Spores 12-15 × 7-8µm. Paraphyses yellow, only slightly swollen from 2µm to 3-4µm at apex. | |||
| Ascophanus bresadolae | ||||
| 27(25) |
|
|||
| - | Spores larger. | 28 | ||
| 28(27) |
|
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| - | Spores shorter. | 29 | ||
| 29(28) |
|
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| - | Spores 13-17 × 7-11µm. | 30 | ||
| 30(29) | Disc punctate with asci. Paraphysis tips swollen up to 3-5µm. Spores 14.5-16 × 9.5-11µm. | |||
| Fimaria leporum | ||||
| - | Disc not punctate with asci. Paraphysis tips not or only slightly swollen. | 31 | ||
| 31(30) | Apothecia pale yellowish. Spores 13-15.5 × 7.5-8.5µm. | |||
| Fimaria theioleuca | ||||
| - | Apothecia chestnut/purplish brown. Spores 14-17 × 7-8.5µm. | |||
| Fimaria cervaria | ||||
| 32(24) | Spores less than 10µm long. | 33 |
| - | Spores mostly longer than 10µm. | 36 |
| 33(32) | Paraphyses markedly capitate to 5-6µm, with yellowish green contents. Apothecia dull at first, yellowish at maturity. Spores 7-10 × 2-4.5µm. | |
| Thelebolus microsporus (fig. 5) | ||
| - | Paraphyses only slightly inflated above, without coloured contents. Apothecia whitish or grey. | 34 |
| 34(33) | Spores 5-7 × 3-4µm. Asci 38-42 × 6-7µm. Apothecia smoky grey, 0.3-0.4mm diam. | |
| Ascophanus cinerellus | ||
| - | Spores larger. Apothecia pale, white or yellowish. | 35 |
| 35(34) | Apothecia up to 1.2mm diam. Asci short stalked, 40-55 × 8-12µm. Spores 7.5-9 × 4.5-5.5µm. | |
| Coprotus glaucellus | ||
| - | Apothecia 0.2-0.5mm diam. Asci attenuate below, 65-85 × 10-15µm. Spores 8-10 × 5-6.5µm. | |
| Coprotus lacteus | ||
| 36(32) | Apothecia chestnut brown up to 1mm diam. Asci 160 × 13µm. Spores 13-16 x 8-11µm. Paraphyses forked, with swollen tips. | |
| Ascophanus misturae | ||
| - | Apothecia lighter coloured. Asci less than 150µm long. | 37 |
| 37(36) | Spores 14-18 × 9-11µm. Apothecia pale yellow/orange, up to 1.5mm diam. Asci cylindrical, 110-150 × 12-15µm. Paraphyses yellowish, slightly inflated to 4-5µm at apices. | |
| Coprotus ochraceus | ||
| - | Spores less than 15µm long. Apothecia up to 0.6mm diam. Asci less than 100µm long. | 38 |