Keys to Fungi on Dung by Watling, Roy

Transcriber's note: Text enclosed by underscores is in italics (_italics_). 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

INTRODUCTION

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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Benjamin, R. K. (1965). Addenda to the merosporangiferous Mucorales. _Aliso_ 6, 1-10. (The 4 papers above are an excellent account of _Syncephalis_, _Piptocephalis_, _Coemansia_ and other unusual allied phycomycetes, republished (1967) as _Bibliotheca Mycologica_ 5 by J. Cramer, Lehre).

Gams, W. & Moreau, R. (1959). Le genre _Mortierella_. _Annales scientifiques de l'Université de Besançon_, Series 2 3, 95-105.

Hesseltine, C. W. (1955). Genera of Mucorales with a note on their synonymy. _Mycologia_ 47, 344-363. (With good key; many other papers by Hesseltine, with others, in _Mycologia_, _American Journal of Botany_, _American Midland Naturalist_ and _Lloydia_).

Ingold, C. T. & Zoberi, M. H. (1963). The asexual apparatus of Mucorales in relation to spore liberation. _Transactions of the British Mycological Society_ 46, 115-134.

Naumov, N. A. (1939). Clés des Mucorinées. _Encyclopédie mycologique_ 9, 1-137.

Zycha, H., Siepmann, R. & Linneman, G. (1969). _Mucorales._ J. Cramer, Lehre. (A revision of Zycha, 1935).

GENERAL REFERENCES

Bell, A. (1983). _Dung Fungi: an illustrated guide to coprophilous fungi in New Zealand._ Victoria University Press, Wellington.

Bon, M. (1987). _The Mushrooms and Toadstools of Britain and North-western Europe._ Hodder & Stoughton, London.

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.

Domsch, K. H., Gams, W. & Anderson, T. H. (1980). _Compendium of soil fungi._ Academic Press, New York.

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.

Key 1. Ascomycota

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_

[Illustration: FIG. 1. _Cheilymenia stercorea_, apothecium.]

[Illustration: FIG. 2. _C. stercorea_, stellate and rooted hairs.]

[Illustration: FIG. 3. _Lasiobolus ciliatus_, apothecium.]

[Illustration: 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) Apothecia brownish, large, 1cm diam. or more. (_Peziza_) 21 - 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 - Spores not apiculate, 20-22 × 8-10µm. _Thecotheus agranulosus_

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 - Apothecia discrete, brownish or purple. (_Fimaria_) 27

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 8-9.5 × 4-4.5µm. _Fimaria equina_ - Spores larger. 28

28(27) Spores 20-38 × 10-13µm. _Fimaria hepatica_ - Spores shorter. 29

29(28) Spores 10-13 × 7-9µm. _Fimaria porcina_ - 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

[Illustration: FIG. 5. _Thelebolus microsporus_, ascus and paraphysis.]

[Illustration: FIG. 6. _Ascodesmis microscopica_, ascospores.]

38(37) Apothecia bright yellow. Asci cylindrical clavate, attenuate below, 65-90 × 10-15µm. Spores 12-14 × 6-8.5µm. Paraphyses branched, apices inflated to 4-5µm, with yellow contents. _Coprotus aurorus_ - Apothecia white/pale yellow, with darker margin. Asci broadly clavate, stalked below 40-55 × 15-30µm. Spores 9-15 × 6.5-9.5µm. Paraphyses inflated above to 5-8µm, hyaline. _Coprotus granuliformis_

39(5) Spores spherical or broadly ellipsoid, brown, ornamented with warts, anastomosing ridges or a reticulum. Asci clavate. Apothecium without excipulum. (_Ascodesmis_, fig. 6) 40 - Spores ellipsoid or spherical, hyaline at first, then purple, becoming brown at maturity; epispore smooth, finely verruculose, warted or cracked. Asci cylindrical. Excipulum present. 45

40(39) Spores 18-21.5 × 13.5-17.5µm. _Ascodesmis macrospora_ - Spores up to 16µm. 41

41(40) Spores ± spherical, L/B ratio mostly up to 1.2. 42 - Spores ± broadly ellipsoidal, L/B ratio mostly 1.2 or more. 43

42(41) Spores ornamented with round warts, 8.5-11 × 8.3-10µm. _Ascodesmis nana_ - Spores ornamented with a network of ridges, 10.5-14 × 9-12µm. _Ascodesmis sphaerospora_

43(41) Spores with a prominent reticulum of ridges (fig. 6), 11-15.5 × 8-13.5µm. Apothecia 150-300µm diam. _Ascodesmis microscopica_ (fig. 6) - Spore ornament not a reticulum. 44

44(43) Spores with 1 simple or branched ridge and isolated or occasionally connected warts, 11-14.5 × 7-11.5µm. Apothecia up to 500µm diam. _Ascodesmis porcina_ - Spores with isolated warts, some joined to form short ridges, but not a reticulum, often capitate, 9.5-12.5 × 7.5-10µm. Apothecia 50-150µm diam. _Ascodesmis nigricans_

45(39) Spores separate in the ascus. (_Ascobolus_) 46 - Spores firmly joined together, both in the ascus and after ejection (fig. 10). (_Saccabolus_) 66

46(45) Spores spherical. 47 - Spores ellipsoid. 48

47(46) Spores 10.5-13.5µm, epispore with numerous but isolated warts. _Ascobolus brassicae_ (figs 8, 9) - Spores 11.5-13.5(15)µm, epispore with subparallel occasionally anastomosing lines. _Ascobolus crosslandii_

48(46) Spores very large, mostly 50-70 × 25-35µm, almost oblong with rounded ends, typically with few cracks in the epispore. _Ascobolus immersus_ (figs 7, 9) - Spores smaller, with epispore smooth, warted or with cracks. 49

49(48) Epispore strongly and irregularly wrinkled with a vesiculose layer of pigment, 11.6-16 × 6.5-9.3µm. Paraphyses capitate up to 18µm. Apothecia up to 0.6mm diam. _Ascobolus rhytidiosporus_ - Epispore not strongly wrinkled/vesiculose. 50

50(49) Epispore basically smooth or warted, perhaps with a few irregular cracks. 51 - Epispore with a clear pattern of cracks or lines. 56

[Illustration: FIG. 7. Apothecia of, from left, _Ascobolus furfuraceus_, _A. immersus_ and _A. albidus_.]

[Illustration: FIG. 8. _A. brassicae_, ascus with spores and detail of operculum.]

[Illustration: FIG. 9. Ascospores of, clockwise from left, _A. immersus_, _A. stictoideus_, _A. albidus_, _A. brassicae_ and _A. crenulatus_.]

51(50) Spores up to 25µm long. 52 - Spores longer, 25µm or more. 54

52(51) Epispore smooth, finely granular or punctate. Gelatinous material unilateral, not surrounding spore. 53 - Epispore warted, spores 18.5-21(22.5) × (9)10-11.5µm, surrounded by gelatinous sheath. _Ascobolus hawaiiensis_

53(52) Spores 18-24 × 10-13µm. Hymenial mucus greenish yellow. Excipulum not brown. _Ascobolus mancus_ - Spores 20-25 × 11-13µm. Hymenial mucus sulphur yellow. Excipulum with rich brown intercellular pigment. _Ascobolus boudieri_

54(51) Epispore smooth or finely granular, spores 23-29(32) × 12-17µm. _Ascobolus elegans_ - Epispore warted. 55

55(54) Spores with a regular pattern of warts and intact epispore, 26-32 × 15-17.5µm. _Ascobolus stictoideus_ (fig. 9) - Spores with irregular patches of thicker pigment, especially at the poles, 28-35 × 16-18µm. _Ascobolus degluptus_

56(50) Spores mostly 18 × 10µm or larger. 57 - Spores mostly smaller than 20 × 10µm. 61

57(56) Apothecia small, mostly up to 1mm diam., colourless. Spores 20-35 × 11-14µm, epispore cracks distant, irregular, often anastomosing. _Ascobolus albidus_ (figs 7, 9) - Apothecia larger, usually 1mm diam. or more, disc yellowish, greenish, purplish or brownish. 58

58(57) Apothecia crowded, purplish or purplish brown with intercellular pigment. Spores 18-28 × 10-12µm, with longitudinal anastomosing cracks. _Ascobolus roseopurpurascens_

- Apothecia yellowish or greenish. 59

59(58) Spores 17-22 × 9.5-12µm with a few widely spaced and irregularly oriented cracks. _Ascobolus michaudii_ - Spores with closely spaced, ± longitudinal, cracks, with varying degrees of anastomosis. 60

60(59) Apothecia furfuraceous, sessile. Ascus wall blue in iodine. Spores 19-28 × 10-14µm. _Ascobolus furfuraceus_ (fig. 7) - Apothecia smooth, substipitate. Ascus wall only faintly blue in iodine. Spores 19-22 × 9.5-13µm. _Ascobolus perplexans_

61(56) Apothecia large, stipitate, 5-10mm diam. Spores 16-19.5 × 8.5-10µm, with subparallel, longitudinal, only rarely anastomosing lines. _Ascobolus lignatilis_ - Apothecia up to 2mm diam. 62

62(61) Apothecia white. 63 - Apothecia yellow, green or brownish. 64

63(62) Spores 13-17 × 7.5-8.5µm, with a coarse reticulum of fine cracks when mature. Only recorded on grouse, capercaillie etc. (Tetraonidae) dung. _Ascobolus carletonii_ - Spores 16-20 × 8-10µm, with a pattern of longitudinal anastomosing cracks. Only recorded on deer dung. _Ascobolus sacchariferus_

64(62) Spores 14.5-16 × 8-9µm, epispore lines not densely crowded. _Ascobolus cervinus_ - Spores smaller, epispore with densely crowded, rarely anastomosing cracks. 65

65(64) Apothecia greenish yellow, furfuraceous, with crenulate margin. Spores 9.5-15 × 6-8µm. _Ascobolus crenulatus_ (fig. 9) - Apothecia brownish yellow to brown, smooth, with undifferentiated margin. Spores 12.5-14.5 × 7-8.5µm. _Ascobolus minutus_

66(45) Asci 4-spored. Spore clusters 42-58 × 14-20µm. Spores 16.5-23 × 9.5-12µm, smooth to finely punctate, but with a thick cap or girdle of reticulated or warted pigment. _Saccobolus quadrisporus_ - Asci 8-spored. 67

67(66) Spore clusters ± globular, 17-26(39) × 15-20µm. 68 - Spore clusters elongated, 2-3 times as long as wide. 69

68(67) Spore clusters compact, subglobose, with only the exposed surface of spores pigmented, ornamented with small and coarse warts. _Saccobolus dilutellus_ - Spores loosely united in cluster, ornamented with small isolated warts covering most of their surface. _Saccobolus globuliferellus_

69(67) Apothecia yellow. Spores in 4 rows of 2 longitudinally arranged spores (fig. 10). 70 - Apothecia hyaline or violaceous (some mature darker). Spores in 2 rows of 3 and 1 row of 2 (fig. 10). 73

70(69) Spore clusters 40µm or longer. 71 - Spore clusters up to 40µm long. 72

71(70) Spore clusters 50-71 × 16-25µm. Spores 22-29 × 8.5-14.5µm, smooth or rarely finely punctate, with distant irregular cracks. _Saccobolus glaber_ (fig. 10) - Spore clusters 43-51 × 14-17µm. Spores 16-22 × 7.5-9µm, with fine isolated warts. _Saccobolus citrinus_

72(70) Spores 14-17.5(19.5) × 7.5-8.5(10)µm, easily separated at maturity. Spore clusters becoming shorter and more rounded with maturity. Apothecia up to 300µm diam., inconspicuous due to their solitary nature and the predominantly brownish colour due to the mature spores. _Saccobolus truncatus_ (fig. 10) - Spores 11.5-13.5 × 5.5-6.5µm. _Saccobolus minimus_

73(69) Apothecia white, covered with tapering squamules composed of septate hyphae. Spore clusters 38-43 × 15-17µm. Spores 16-17.5 × 7-8.5µm, smooth or finely punctate. _Saccobolus caesariatus_ - Apothecia not white, without tapering scales. 74

74(73) Spore clusters mostly over 40µm long. 75 - Spore clusters mostly under 40µm long. 76

75(74) Spore clusters 38-62 × 14-19µm. Spores 13-21.5 × 6.5-9.5µm, smooth, finely warted or with reticulate cracks. Apothecia 0.2-2mm diam. _Saccobolus versicolor_ (fig. 10) - Spore clusters 42-60 × 18-24µm. Spores very coarsely warted, 17.5-23 × 8.5-10µm (inc. warts). _Saccobolus beckii_

[Illustration: FIG. 10. Spore clusters of, from left, _Saccobolus versicolor_, _S. glaber_ and _S. truncatus_.]

76(74) Spore clusters compact, 26-43 × 13-19µm. Spores 13.5-18 × 7.5-9.5µm, epispore with fine or coarse warts. Apothecia 0.3-0.8mm diam. _Saccobolus obscurus_ - Spore clusters elongated, 28-37 × 10-13µm. Spores 10-14.5 × 5-7.5µm, epispore smooth or very finely granular. Apothecia 0.1-0.3mm diam. _Saccobolus depauperatus_

77(4) Asci operculate or bursting, without a subapical ring. Spores ellipsoid. 78 - Apothecia white, often minutely hairy at the margin. Ascus dehiscing by a vertical slit; the slit is prevented from running right down the ascus by a subapical ring of thickening. Spores ellipsoid-fusiform. (_Ascozonus_, figs 14, 15) 90

78(77) Asci 16-spored. Spores ellipsoid, 11-16 × 7-10µm. _Coprotus sexdecemsporus_ - Asci more than 16-spored. 79

79(78) Asci 32-spored. 80 - Asci more than 32-spored. 84

80(79) Asci very large, nearly 0.5mm long, spores 30-35 × 13-17µm (32-40 × 20-24µm in Kimbrough, 1969). Apothecia pale coloured. _Thecotheus pelletieri_ - Asci and spores smaller. 81

81(80) Spores 10µm or longer. 83 - Spores up to 10µm long. 82

82(81) Spores ellipsoid, with minute scattered warts visible under oil-immersion, 7-9 × 4-4.5µm. Apothecia densely crowded, 90-120µm diam., with 8-13 asci. Asci 32-55 × 16-18µm with (24-)32 spores. Paraphyses 1.5-2µm, clavate to 4-4.5µm. _Thelebolus caninus_ - Spores subacute at apices, ca 6 × 4µm (described as 'minute'; this value is suggested by Boudier's comparison with _R. dubius_, for which measurements are given). Apothecia densely crowded, tawny yellowish-brown. _Ryparobius brunneus_

83(81) Spores 10-12.5 × 5-7.5µm. Asci clavate, 75-100 × 20-30µm. Paraphyses enlarged to 6µm at apex. _Coprotus albidus_ - Spores 13.5-17.5 × 7-8µm. Asci 10-15 per apothecium, 120-175 × 50-75µm. Paraphyses filiform. _Coprotus rhyparobioides_

84(79) Asci with up to 64 spores. 85 - Asci with many more than 64 spores--impractical to count. 86

85(84) Asci 64-spored, broad clavate with short stalk, 80-130 × 30-60µm. Spores 8-12 × 4-7µm. _Coprotus niveus_ - Asci broadly clavate with up to 64 spores, 60-100 × 20-30µm. Spores 7-10 × 4.5-5.5µm. Apothecia superficial, on the surface of the substrate, yellowish brown, gregarious, united into a crust. _Thelebolus crustaceus_

86(84) Apothecia superficial, 400-600µm diam., with prominent, acuminate, superficial, 1-2-septate hairs, 80-190µm long, often roughened towards their apex, with one 1000+-spored ascus, 110-240 × 15-27µm. Spores very variable, 6.5-16 × 3.7-8.8µm (mostly 7.5-13 × 4.5-7µm). _Lasiobolus monascus_ - Apothecia minute, rarely above 350µm diam., globose and immersed in substrate when young. Asci broad globose, with 100-200 spores. Usually only 1-3 asci in each apothecium, which dehisce by bursting at the apex. 87 (Other _Ryparobius_ spp. will key out here [e.g. _R. dubius_, _R. myriosporus_, _R. pachyascus_ and _R. polysporus_]. They all have scattered to gregarious, immersed to semi-immersed apothecia 100-200µm diam., with relatively few asci, each with 100-250 ellipsoid to subacuminate ca 5-7 × 3-4µm spores. There are insufficient modern observations to allow their identification and separation with confidence).

87(86) Apothecia with a few, but obvious, setae. Spores 9 × 7µm or larger. 88 - Apothecia without setae. Spores ellipsoid, 6-9 × 3.5-4µm. 89

88(87) Spores ellipsoid, 9-11 × 7-9µm. Setae up to 600µm long. _Trichobolus zukalii_ - Spores subglobose, 11-12 × 10-11µm. Setae up to 300µm long. _Trichobolus sphaerosporus_ (fig. 11)

89(87) Apothecia and asci large, 170-250µm diam. _Thelebolus stercoreus_ (fig. 12) - Apothecia and asci small, rarely above 80-90µm diam. _Thelebolus nanus_ (fig. 13)

90(77) Asci 16(-24)-spored. Spores not closely aggregated into an imbricated mass, 13-14 × 6µm (8-9 × 4µm)[1]. Apothecial hairs rough, subulate. _Ascozonus parvisporus_ - Asci with 32 or more spores. 91

91(90) Asci 32-spored. Spores 16.5-18 × 4.5-5µm (11-12 × 3-3.5µm)[1]. Apothecia with a single row of sharp, pointed, roughened hairs. _Ascozonus crouanii_ - Asci more than 32-spored. 92

92(91) Asci 48-spored. Spores spindle-shaped, 12-14.5 × 2.5-4µm. _Ascozonus leveillei_ - Asci more than 48-spored. 93

93(92) Asci 64-spored. 94 - Asci more than 64-spored. 95

94(93) Apothecia with a short base of globose cells, with minutely roughened marginal hairs up to 30 × 8µm. Spores elliptic-fusoid, 12-14 × 3-5µm. _Ascozonus woolhopensis_ (figs 14, 15) - Apothecia sessile, with aseptate smooth hairs. Spores 21 × 7.5µm (13-14 × 4.5-5µm)[1]. _Ascozonus cunicularis_

[Illustration: FIG. 11. _Trichobolus sphaerosporus_, apothecium.]

[Illustration: FIG. 12. _Thelebolus stercoreus_, apothecium.]

[Illustration: FIG. 13. _T. nanus_, mature and immature apothecia, and detail of ascus dehiscence.]

[Illustration: FIG. 14. _Ascozonus woolhopensis_, apothecium and apothecial hair.]

[Illustration: FIG. 15. _A. woolhopensis_, ascus with spores and detail of dehiscence.]

95(93) Apothecia with a short base of globose cells, with short, irregular hairs. Asci 64-96-spored Spores elliptic-fusoid, 14-14.5 × 5-5.5µm (10-15 × 3.5-4µm)[1]. _Ascozonus leveillanus_ - Apothecia sessile, dotted with hairs in connate groups of 2-3. Asci with 128 or more spores. Spores 10 × 5µm (7 × 3.5µm)[1]. _Ascozonus subhirtus_

96(3) Apothecia stalked. 97 - Apothecia not stalked. 98

97(96) Apothecia up to 2mm diam., with a short cylindrical stalk, light brown. Asci 150 × 10µm. Spores hyaline, with 2 oil drops, occasionally 1-septate, 13-15 × 4.5µm. _Lanzia cuniculi_ - Apothecia up to 3mm diam., pale olivaceous to grey, with a long, slender, reddish-brown stalk arising from a sclerotium in the dung. Asci 30-40 × 4-5µm. Spores ellipsoid, grey-brown, 4-4.5 × 2µm. _Martininia panamaensis_

98(96) Spores 7-11(14) × 1.75-2.75µm. ellipsoid, ellipsoid- fusiform or slightly clavate. Apothecia yellowish brown when fresh, drying darker, up to 1mm diam. Asci 42-60 × 7.5-9µm, pore weakly blue in iodine. _Pezizella albula_ - Spores and asci smaller. 99

99(98) Spores linear, 3-5 × 1µm. Asci 30 × 5µm, cylindrical with a short stipe. Paraphyses not clavate but fused to form an epithecium. Apothecia pale pellucid, 0.5-1mm diam. _Orbilia leporina_ - Spores longer, subulate, curved. 100

100(99) Spores 7-8.5 × 1.2-1.8µm. Asci 36-40 × 3-5µm, gradually tapering to a short base. Paraphyses enlarged to 3µm at apex, covered with brown granules. Apothecia light brown, 0.4-1.mm diam. _Orbilia fimicola_ - Spores 8-10.55 × 0.9-1µm. Asci 30-45 × 3µm, cylindrical-clavate with narrow tapering base and truncate apex. Paraphyses 2µm diam., the tips with a crust-like secretion fusing together to form a shiny epithecium. Apothecia white to yellowish, 180-700µm diam. _Orbilia fimicoloides_

Key 2. Perithecial, pseudothecial, cleistothecial and gymnothecial fungi

1 Perithecia occurring singly or in groups, but directly (key 1,2) on the dung or buried in it (figs 16, 28, 19, 22, 27, 30, 32, 34-36). 2 - Perithecia occurring in or on a mass of fungal tissue (stroma) growing in or on the dung (figs 32, 37). 135

2(1) Spores black, brown or dark olive-greenish. 3 - Spores hyaline or pale coloured, at least under the microscope (may be coppery red _en masse_). 117

3(2) Spores smooth, without an ornamentation of hyaline pits. 4 - Spores 1-celled, ornamented with hyaline pits. (_Gelasinospora_) 114

4(3) Perithecia dark, olive, brown or black. 5 - Perithecia reddish brown, orange or golden, globose, with a neck. Spores black, limoniform. 116

5(4) Perithecia globose, surmounted by a dense tuft of greyish green hairs, which may be branched or simple, straight or curly. Spores olivaceous, limoniform. Asci clavate, soon disappearing. (A large genus not characteristic of dung, but occurring occasionally). _Chaetomium_ (fig. 16) - Perithecia more pyriform, or if globose then with a distinct neck, may be setose but not densely hairy, with clavate or cylindrical asci. 6

6(5) Each spore composed of 4 or more cells in a row (figs 17, 21). Asci bitunicate (figs 20, 23). 7 - Spores 1- or 2-celled. Asci bitunicate or unitunicate. 29

7(6) Spores 16-32-celled, united firmly together in a bundle both in the ascus and after discharge. Germ slits usually absent. (_Sporormia_) 8 - Spores each with 4 or more cells, each spore free and surrounded by its own gelatinous sheath. Germ slits usually present. (_Sporormiella_) 11

8(7) Spores 16-20-celled. 9 - Spores 29-32-celled, 130-160 × 4-6µm. _Sporormia mirabilis_

9(8) Spores 16-celled, 85-116 × 5-6.5µm. _Sporormia fimicola_ - Spores smaller. 10

10(9) Spores 16-celled, 37-45 × 3µm. Asci 50-60 × 10-12µm. _Sporormia sp._ (fig. 17) [recorded as _S. fimetaria_ by Richardson (1972); see also Bell (1983) and Dissing (1992)] - Spores 16-20-celled, 50-57 × 3.5-4.5µm. Asci 70-80 × 12-16µm. _Sporormia fimetaria_ (These two taxa may represent the extremes of _S. fimetaria_).

11(7) Spores 4-celled. 12 - Spores more than 4-celled. 22

12(11) Spores more than 65-70µm long. 13 - Spores less than 65-70µm long. 15

[Illustration: FIG. 16. _Chaetomium_ sp., perithecium and spore.]

[Illustration: FIG. 17. _Sporormia_ sp., ascus and spores.]

[Illustration: FIG. 18. _Sporormiella ovina_, pseudothecium.]

[Illustration: FIG. 19. _S. intermedia_, pseudothecium.]

[Illustration: FIG. 20. _S. intermedia_, immature bitunicate ascus and mature ascus with outer layer ruptured.]

[Illustration: FIG. 21. Ascospores of, from left, _S. ovina_, _S. intermedia_ (with gelatinous sheath characteristic of the genus), _S. lageniformis_, _S. vexans_, _S. bipartis_ and _S. minima_.]

13(12) Spores 65-95 × 15-18µm. _Sporormiella megalospora_ - Spores longer than 90µm. 14

14(13) Spores 90-118 × 15-20µm. Asci tapering gradually from the broadest part near the apex to a 'stipe'. _Sporormiella ovina_ (figs 18, 21) - Spores 91-114 × (14)18-21µm. Asci cylindrical, abruptly contracted below to a short 'stipe'. _Sporormiella borealis_

15(12) Spores mostly less than 35µm long. 16 - Spores mostly between 35-60µm long. 19

16(15) Spores less than 25µm long. 17 - Spores 25-35(38)µm long. 18

17(16) Spores (15)17-24(26) × 5-7µm, end cells broadly conical. Ascospores uniseriate. Asci 120-135µm long. Pseudothecia 250-300µm diam. _Sporormiella pulchella_ - Spores 16-22 × 4.5-5.5µm, end cells subovate. Ascospores biseriate. Asci 95-125µm long. Pseudothecia 300-350µm diam. _Sporormiella nigropurpurea_

18(16) Spores 30-38.5 × 5.5-6.5µm. Asci clavate, tapering gradually below to a 'stipe'. _Sporormiella leporina_ - Spores 27-36(38) × 4-6(8)µm, tending to break in two at the middle septum. Asci cylindrical, abruptly contracted below. _Sporormiella minima_ (fig. 21)

19(15) Spores with end cells rounded. Asci cylindrical, abruptly contracted below. 20 - Spores with end cells tapered and slightly conical. Asci clavate, tapering gradually to a long stalk. 21

20(19) Spores 45-65 × 8-11.5µm. _Sporormiella intermedia_ (figs 19-21) - Spores 38-46 × 6.5-8µm. _Sporormiella australis_

21(19) Spores 45-60 × 11.5-14µm, germ slits parallel with long axis. _Sporormiella grandispora_ - Spores 35-45(48) × 7-9(10)µm. _Sporormiella lageniformis_ (fig. 21)

22(11) Spores 5-celled, 70-80 × 17-19µm. _Sporormiella pentamera_ - Spores more than 5-celled. 23

23(22) Spores 7- or 8-celled. 24 - Spores 13-celled, 46-60 × 9-10µm. _Sporormiella antarctica_

24(23) Spores 7-celled. 25 - Spores 8-celled. 26

25(24) Spores 40-55 × 7-9µm, readily disarticulating, the end cells longer than wide, the rest shorter than wide. _Sporormiella vexans_ (fig. 21) - Spores 70-80 × 16-18µm, end cells rounded. _Sporormiella heptamera_

26(24) Spores mostly longer than 45µm. 27 - Spores less than 50µm long, not disarticulating at the central septum. 28

27(26) Spores 45-60 × 5-7.5µm, disarticulating at the central septum, all cells the same width. _Sporormiella bipartis_ (fig. 21) - Spores 50-59 × 10-12µm, not disarticulating, 3rd cell down wider than the others. _Sporormiella corynespora_

28(26) Spores (33)37-40(49) × 7-9µm, cylindrical. Asci abruptly contracted below. _Sporormiella pascua_ - Spores 40-48 × 7-8µm, fusiform cylindrical. Asci gradually tapered below. _Sporormiella octomera_

29(6) Spores obviously 2-celled at maturity. 30 - Spores 1-celled, or appearing 1-celled at maturity. (Those of _Podospora_, _Schizothecium_ etc. are 2-celled in early stages of their development, but only one cell matures to become pigmented; the other remains hyaline, often collapses, and may be difficult to see). 47

30(29) Spores 23-28 × 13-17µm, upper cell dark, 15-19µm, with close, blunt spines giving the impression of a pitted spore surface, with apical germ pore, the lower cell hyaline, 6-8.5µm, smoky-brown. Asci unitunicate, 4-spored. Perithecia 400µm diam. _Apiosordaria verruculosa_ (fig. 24) - Both cells of spore similar in shape, size and colour. 31

31(30) Asci unitunicate. Spores with a 'gelatinous' appendage at each end. Perithecial neck with setae. 32 - Asci bitunicate. Spores without gelatinous appendages, although a sheath may be present. 33

32(31) Spores 38-48 × 11-14µm, appendages longitudinally fibrillate. _Zygospermella striata_ - Spores 46-68 × 11-17µm, appendages hollow, not fibrillate. _Zygospermella insignis_ (fig. 25)

33(31) Spores with each end truncated by a germ pore. Pseudothecia with dark bristles at neck. (_Trichodelitschia_) 34 - Spores with rounded ends and germ slits along the sides. Pseudothecial neck smooth or hairy, but without setae. (_Delitschia_, fig. 26) 36

34(33) Spores 28-34 × 9-12µm. _Trichodelitschia aedelphica_ - Spores smaller. 35

35(34) Spores 20-27.5 × 8-11µm. _Trichodelitschia bisporula_ (figs 22, 23) - Spores 18-21 × 6-7µm. _Trichodelitschia munkii_

36(33) Asci ca 256-spored. Spores 14-15 × 6-8µm. _Delitschia myriaspora_ - Asci 8-spored. 37

37(36) Spores less than 20µm long. 38 - Spores more than 20µm long. 41

38(37) Spores 8-11 × 3-5µm. _Delitschia perpusilla_ - Spores 10-20µm long. 39

39(38) Spores 10-14 × 5-6µm. _Delitschia marchalii_ - Spores longer. 40

40(39) Spores 14-18 × 6-10µm, uniseriate. Asci 70-90 × 7-16µm. _Delitschia niesslii_ - Spores (16)18-20(22.5) × 6-7.5µm, biseriate. Asci 80-145 × 20-25µm. _Delitschia consociata_ (fig. 26)

41(37) Spores mostly wider than 20µm. 42 - Spores mostly less than 20µm wide. 43

42(41) Spores 50-64 × 19-23µm. _Delitschia furfuracea_ - Spores 50-70 × 25-33µm. _Delitschia winteri_ (fig. 26)

43(41) Spores 20-25 × 4.5-6µm, the cells slightly tapered and almost completely separated. Pseudothecia hairless, globose, ca 200µm diam. _Delitschia leptospora_ (fig. 26) - Spores longer and wider. 44

44(43) Spores transversely septate. 45 - Spores obliquely septate, deeply constricted at the septum, 35-50 × 15-18µm. _Delitschia didyma_

45(44) Pseudothecia hairy. Spores 37-50 × 17-20µm, not deeply constricted at the septum. _Delitschia chaetomioides_ - Pseudothecia smooth. 46

46(45) Spores biseriate, 45-55 × 13-16µm, one cell usually larger than the other, deeply constricted at the septum and readily separating. _Delitschia canina_ - Spores uniseriate, 40-55 × 16-21µm, both cells equal. _Delitschia patagonica_

[Illustration: FIG. 22. _Trichodelitschia bisporula_, pseudothecium.]

[Illustration: FIG. 23. _T. bisporula_, expanded ascus broken through the outer wall, with spores.]

[Illustration: FIG. 24. _Apiosordaria verruculosa_, ascospores.]

[Illustration: FIG. 25. _Zygospermella insignis_, ascus and ascospore.]

[Illustration: FIG. 26. Ascospores of, from left, _Delitschia winteri_, _D. consociata_ and _D. leptospora_.]

47(29) Spores with colourless 'gelatinous' secondary appendages (caudae, fig. 28) at one or both ends (not always easy to see; mounting in Indian ink is useful, and essential for some). A hyaline (empty) cell, the primary appendage (fig. 28), may also be present. 48 - Spores without caudae, although a colourless gelatinous sheath may be present. Primary appendages present or absent. 88

48(47) Perithecia often hairy or tomentose when young. Immature spores long, wavy cylindrical, with a row of globules, and more likely to be seen than mature spores (fig. 29). Secondary appendages thin, simple, up to 60 × 3µm. Mature spores with a dark cell 14-25 × 7-13µm and pedicel (primary appendage) 25-50 × 3-6µm. (_Cercophora_) 49 - Perithecia often with scales or setae at the neck or tomentose. Caudae, simple or compound. Immature spores clavate or ellipsoid, not long, wavy cylindrical. Mature spores readily observed. 51

49(48) Immature spores 45-70 × 4-6µm. 50 - Immature spores smaller, 38-52 × 3-3.5µm. Mature spores with upper (dark) cell 14-18 × 7-9µm; hyaline pedicel 27-36 × 3-3.5µm. _Cercophora silvatica_

50(49) Perithecia with white or grey tomentum. Young spores 45-65 × 4.5-6µm. Mature spores with upper cell 17-25 × 8.5-13µm and pedicel 30-50µm long. _Cercophora coprophila_ (fig. 29) - Perithecia with flexuose brown hairs and, at the neck, tufts of agglutinated, swollen, obtuse hairs. Young spores 52-68 × 4-5µm. Mature spores with upper cell 15-25 × 9-11µm and pedicel 35-45µm long. _Cercophora mirabilis_

51(48) Primary appendage absent. (_Arnium_, fig. 28) 52 - Primary appendage present. 60

52(51) Asci (64-)128-spored. Spores 18-26 × (10)12-15µm. Perithecial neck sometimes with rigid, brown, septate hairs up to 330µm. _Arnium leporinum_ - Asci 4- or 8-spored. 53

53(52) Asci 4-spored. 54 - Asci 8-spored. 55

54(53) Spores ellipsoid, sometimes inequilaterally flattened, 44-54 × 22-30µm, with 1 apical germ pore, caudae not swelling in water. Perithecium usually with lateral tufts of agglutinated hairs up to 550µm long. _Arnium arizonense_ - Spores evenly ellipsoid-fusiform, 31-55 × 18-25µm, with germ pore at each end, caudae covering germ pores, 35-60 × 7-11µm, but rupturing and swelling to up to 130 × 50µm, and becoming diffuse and irregular. Perithecial neck covered with rigid hairs up to 190 × 2.5µm. _Arnium hirtum_

55(53) Perithecial neck distinctly setose with rigid hairs. 56 - Perithecial neck without setae. 57

56(55) Spores evenly ellipsoid-fusiform, 31-55 × 18-25µm, with germ pore at each end, caudae covering germ pores, 35-60 × 7-11µm, but rupturing and swelling up to 130 × 50µm, and becoming diffuse and irregular. Perithecial neck covered with rigid hairs up to 190 × 2.5µm. _Arnium hirtum_ - Spores slightly inequilateral, 35-43 × 17-23µm, caudae 50-75 × 5-8µm, not covering germ pores. Perithecial neck with brown hairs up to 250µm long. _Arnium cervinum_

57(55) Perithecia covered with a dense tomentum of septate flexuous hairs. Spores mostly longer than 45µm. Only occasionally fimicolous. 58 - Perithecia without a tomentum. Spores up to 45µm. 59

58(57) Spores (40)45-54 × 25-35µm, uniseriate. Tomentum pale or grayish. _Arnium olerum_ - Spores 47-70 x 20-30µm, biseriate above. Tomentum olivaceous brown. _Arnium tomentosum_

59(57) Spores somewhat inequilateral, rounded below, pointed above, 31-40 × 18-24µm, caudae 50-120 × 6-10µm, with 1 apical germ pore not covered by cauda. _Arnium caballinum_ - Spores equilateral, 36-44 × 20-23µm, caudae 50-80 × 6-8µm, covering germ pores. _Arnium mendax_

60(51) Perithecia with scales at the neck, composed of inflated and agglutinated cells (fig. 27, _S. conicum_). (_Schizothecium_) 61 - Perithecia setose or hairy at the neck, but not with inflated cells, or neck black but almost hairless. (_Podospora_) 70

61(60) Asci 4-spored. 62 - Asci 8-spored. 63

62(61) Spores 11-14.5 × 6.5-9µm. _Schizothecium nanum_ (fig. 28) - Spores 19-24 × 12-14.5µm. _Schizothecium tetrasporum_

63(61) Spores more than 30µm long. 64 - Spores less than 30µm long. 65

64(63) Perithecia crowned with a fascicle of long agglutinated hairs at the neck, up to 335µm long. Spores 31-40 × 15-25µm, biseriate. _Schizothecium aloides_ - Perithecia with shorter, less remarkable tufts. Spores 30-45 × 19-24µm, ± uniseriate. _Schizothecium glutinans_

65(63) Perithecial neck with rigid setae, as well as agglutinated hairs (which may be greatly reduced). Asci 140-210 × 19-25µm, broadest at the markedly rounded apex. Spores 18-23 × 11-14µm. _Schizothecium pilosum_ - Perithecial neck without rigid setae. Asci broadest in the middle. 66

66(65) Spores mostly over 23µm long. 67 - Spores up to 23µm long. 69

67(66) Spores 22-25(27) × 11-13µm. Scales at neck distinct. _Schizothecium hispidulum_ - Spores wider, 12-19µm 68

68(67) Perithecia 0.5-1mm high, scales at neck usually well developed. Spores (23)26-30 × 12-17µm. _Schizothecium conicum_ (fig. 27) - Perithecia 1-2mm diam., subpyriform, neck velvety with indistinct scales. Spores 24-28 × 15-19µm. _Schizothecium squamulosum_

69(66) Spores 17-23 × 8.5-13.5µm, primary appendage slender cylindrical, 6-8 × 2µm. Perithecia 0.25-0.7mm high, sometimes with poorly developed scales. _Schizothecium vesticola_ (fig. 28) - Spores 11-14 × 6-8µm, primary appendage short, 2µm long, almost triangular. Perithecia 0.3-0.45mm high, with short agglutinated hairs. _Schizothecium cervinum_

70(60) Asci 4-spored. Spores 35-40 × 18-19µm. _Podospora pauciseta_ - Asci with more than 4 spores. 71

[Illustration: FIG. 27. Perithecia, from left, of _Podospora appendiculata_, _Schizothecium conicum_, _P. excentrica_ and _P. decipiens_, with detail of hairs. FIG. 28. Ascospores of, from left, _Podospora excentrica_, _P. appendiculata_, _S. vesticola_, _S. nanum_, _P. decipiens_, _'P. dagobertii'_ and _Arnium_ sp. FIG. 29. _Cercophora coprophila_, immature (l) and mature (r) ascospores.]

71(70) Asci 8-spored. 72 - Asci with more than 8 spores. 82

72(71) Spores more than 45µm long. 73 - Spores less than 45µm long. 74

73(72) Spores 48-60 × 27-31µm, caudae apparently striate. Perithecia superficial, covered with rigid, nonagglutinated hairs up to 120µm. _Podospora fimiseda_ - Spores 50-68 × 22-32µm, caudae apparently segmented, with an intestine-like appearance. Perithecia immersed to superficial, with a long neck, tomentose with long flexuous hairs when young, more or less glabrous when mature. _Podospora intestinacea_

74(72) Perithecia superficial, ovoid to globose, covered with short (up to 100µm), sparse, radiating, hyaline tipped, hairs. Spores 24-31 × 11-15µm, with simple caudae. _Podospora appendiculata_ (figs 27, 28) - Perithecia with base immersed in substrate, pyriform, without such hairs. 75

75(74) Perithecial neck with short tubercular hairs, up to 20µm long. Spores 32-42 × 17-22µm, with a long but withering primary appendage. Caudae in two rings, one inserted near the base of the primary appendage, the other at the spore apex. The individual filaments may be free, but often clump together to form an apparently broad appendage. _Podospora decipiens_ (figs 27, 28) - Perithecial hairs longer. Caudae single or 4 at each end. 76

76(75) Spores with 4 caudae at each end. 77 - Spores with a single cauda at each end. 78

77(76) Spores 40-45 × 22-25µm. _Podospora gwynne-vaughaniae_ - Spores 29-40 × 16-25µm. _Podospora communis_

78(76) Spores less than 30 × 15µm. 79 - Spores larger than 30 × 15µm. 80

79(78) Spores 21-28 × 11-14µm, primary appendage 12-14 × 4µm. Perithecia 0.3-0.5mm diam., neck setose with rigid cylindrical hairs. Asci 200-250 × 22-26µm, broadest in the middle. _Podospora ellisiana_ - Spores 18-23 × 11-14µm, primary appendage 4-8 × 3µm. Perithecia 0.2-0.3mm diam., neck setose with rigid hairs. Asci 140-210 × 19-25µm, broadest at the markedly rounded apex. _Schizothecium pilosum_

80(78) Perithecia ca O.9-1.4mm high × 0.6-0.7(0.85)mm diam., neck not hairy. Spores (29)36-45 × (17.5)22-27µm, caudae ephemeral and difficult to see, even in Indian ink. _Podospora pyriformis_ - Perithecial neck with tufts of rigid hairs. 81

81(80) Perithecia 0.38-0.53mm high × 0.21-0.38mm diam., ± immersed, with hairs at the neck up to 335µm long, grouped in rigid fascicles. Spores slightly flattened on one side, 30-37 × 18-24µm, caudae invisible in water. _Podospora excentrica_ (figs 27, 28) - Perithecia ca 0.8-1.4mm high × 0.4-0.7mm diam., semi-immersed, hairy all over, flexuous below, rigid and pointed at the neck up to 170µm. Spores 33-45 × 22-27µm. _Podospora perplexens_

82(71) Asci 16-32-spored. Perithecial neck with short tubercular hairs. Spores 25-36 × 15-24µm. Caudae in two rings, one inserted at the base of the primary appendage, the other at the spore apex; individual filaments may be separate or clumped to appear as a broad single appendage (cf. _P. decipiens_). _Podospora pleiospora_ - Asci with more than 32 spores. 83

83(82) Perithecia with tufts of rigid hairs at neck. Asci with more than 64 spores. 84 - Perithecia without tufts of rigid hairs. Asci 64-spored. 87

84(83) Spores 14-17 × 9-11µm. Asci 256-spored. Perithecia ca 500µm diam., immersed, except for the neck, which has tapered tufts of hairs up to 300µm. _Podospora curvicolla_ - Spores larger. Perithecia semi-immersed. 85

85(84) Spores (18)20-26 × 12-16µm, caudae of 2-several filaments covered with granules. Asci 512-spored. Perithecia up to 1mm high × 0.95mm diam., neck with rigid but non-agglutinated hairs up to 130µm long. _Podospora granulostriata_ - Caudae simple, without granular appearance. Asci 128-spored. Perithecia not larger than 750µm high × 500µm diam., with rigid, non-agglutinated hairs up to 190µm long at neck. 86

86(85) Spores 17-19 × 10-12µm. _Podospora setosa_ - Spores 19-24 × 11-16µm. _Podospora tarvisina_ (See discussion in Lundqvist (1972) on these last three names)

87(83) Spores 24-34 × 14-19µm, caudae in two rings, one inserted at the base of the primary appendage, the other at the spore apex; individual filaments may be separate or clumped to appear as a broad single appendage (cf. _P. decipiens_/_P. pleiospora_). Perithecia ca 0.6-1.1mm high × 0.4-0.5mm diam., covered with flexuous hairs or rarely smooth. _Podospora myriaspora_ - Spores 15-20 × 10-15µm, caudae small, simple and evanescent. Perithecia 0.4-0.5mm high, covered with long flexuous hairs. _Podospora collapsa_

88(47) Spores with primary appendage. 89 - Spores without primary appendage. 93

89(88) Spores with primary appendage directed towards base of ascus. 90 - Spores with primary appendage directed towards apex of ascus. (_Anopodium_) 91

[Illustration: FIG. 30. Perithecia of, from left, _Coniochaeta ligniaria_, _C. scatigena_ and _C. hansenii_. FIG. 31. Ascospores of _C. scatigena_ (l) and _C. ligniaria_ (r).]

90(89) Spores 34-45 × 19-25µm, without caudae but surrounded by a thin (ca 5µm) gelatinous sheath. Perithecia ca 0.5-0.7mm diam., ± smooth. _Podospora globosa_ - Spores 17-20 × 8-9.5µm, flattened on one side, convex on the other. Perithecia 0.3-0.45µm diam., with distal cells of agglutinated hairs fimbriate. _Podospora fimbriata_

91(89) Perithecia hairy. Spores 27-32 × 16-19µm, appendage 15-18 × 2.5-3µm. _Anopodium ampullaceum_ - Perithecia glabrous. 92

92(91) Spores 28-32 × 16-21µm, appendage 12-15 × 3-3.8µm. _Anopodium epile_ - Spores 30-37 × 16-20µm, appendage 24-27 × 5µm. _'Podospora' dagobertii_ (fig. 28) (The combination in _Anopodium_ has not been made; see Lundqvist, 1964, 1972)

93(88) Spores flattened, disc shaped, with a germ slit around the edge. Perithecial neck with short (up to 120µm) setae. (_Coniochaeta_, figs 30, 31) 94 - Spores ellipsoid. Perithecial neck without setae or with very prominent (up to 950µm) tufts of agglutinated hairs. 99

94(93) Asci with numerous (64-128) spores. 95 - Asci 8-spored. 96

95(94) Spores 6-10 × 5-9 × 4-7µm. Perithecial setae up to 120µm long. _Coniochaeta hansenii_ (fig. 30) - Spores 13-16 × 9.5-13.5 × 5.5-8µm. Perithecial setae up to 35µm long. _Coniochaeta_ sp.

96(94) Spores 7-9 × 6-8 × 5-6µm, slightly flattened. _Coniochaeta leucoplaca_ - Spores larger. 97

97(96) Spores narrowly elliptical in face view (length more than 2 × width), ca 13-18 × 6-9 × 4-6µm. _Coniochaeta saccardoi_ - Spores broadly elliptical to nearly circular in face view (length less than 2 × width). 98

98(97) Spores (9)10-16(20) × 7.5-10(15) × (4)5-8µm. Neck setae 20-50µm long. _Coniochaeta ligniaria_ (figs 30, 31) - Spores (16)17-23 × (10)13-19 × 7.5-10(15)µm. Neck setae 40-80µm long. _Coniochaeta scatigena_ (figs 30, 31)

99(93) Perithecial neck with prominent agglutinated tufts of rigid setae up to 950µm long. Spores 43-54 × 20-29µm, with apical germ pore. A gelatinous sheath which surrounds the whole spore swells in water, and appears fringed at the margin and radially striate. _Arnium macrothecium_ - Perithecial neck without setae. Gelatinous sheaths may be clearly visible around spores, but are not complex in structure. 100

100(99) Spores with germ slit along the side. Ascus with a large and complex plug at the tip staining blue or red in KI (other genera have asci with blue staining ascus tips, but the feature is very pronounced in this genus and is unlikely to be mistaken). Perithecia form singly or severally in a stroma which is usually of limited extent, often without a definite margin. [N.B. if orange and with a stroma see _Selinia_, 119]. (_Hypocopra_, fig. 32) 101 - Spores without germ slits, but often asymmetrical, and with a small papilla at the basal end. Asci without complex apical plug. (_Sordaria_, fig. 33) 107

101(100) Spores mostly less than 25µm long. 102 - Spores more than 25µm long. 104

102(101) Spores 9-14 × 6-7µm. _Hypocopra parvula_ - Spores larger. 103

103(102) Stroma with a brown hyphal mat between perithecial necks. Spores 19-27 × 10-14µm. _Hypocopra equorum_ (fig. 32) - Stroma with white hyphae between black perithecial necks, becoming smooth. Spores 23-25 × 12-14µm. _Hypocopra brefeldii_

104(101) Ascospores up to 15µm wide. 105 - Ascospores 15µm or wider. 106

105(104) Ascospores 25-31 × 10-15µm, distinctly flattened on one side. Ascus plug blue in KI, but becoming reddish. _Hypocopra planispora_ - Ascospores 26-32 × 13-14µm, ellipsoid and narrowed towards their ends. _Hypocopra stephanophora_

106(104) Ascospores 27-43 × 16-20µm. _Hypocopra merdaria_ - Ascospores 38-50 × 19-24µm. _Hypocopra stercoraria_

107(100) Spores up to 10µm long. 108 - Spores 10µm or longer. 109

108(107) Asci 8-spored. Spores 8 × 4µm. _Sordaria minima_ - Asci ca 128-spored. Spores 5-8 × 4-5µm. _Sordaria polyspora_

109(107) Spores relatively narrow, at least twice as long as wide, 22-26 × 9-12µm. Gelatinous sheath broad, distinct. _Sordaria alcina_ - Spores relatively broad, less than twice as long as wide. 110

110(109) Spores mostly 25µm or longer. 111 - Spores up to 25µm long. 112

111(110) Spores (21)23-29(30) × 14.5-17(18)µm, with apiculate base. Gelatinous sheath broad, distinct. Asci 240-300 × 20-24µm. _Sordaria superba_ - Spores (26)28-35 × (17)18-22µm, with slightly apiculate base. Gelatinous sheath broad, distinct. Asci 280-350 × 30-35µm. _Sordaria macrospora_

112(110) Spores with gelatinous sheath absent or very thin, 19.5-25 × 15.5-19µm. _Sordaria humana_ (fig. 33) - Spores with gelatinous sheath, up to 15µm diam. 113

113(112) Spores obovoid to broadly ellipsoid, 18-23 × 12-15µm. _Sordaria lappae_ - Spores ellipsoid, 17-25 × 10-14µm. _Sordaria fimicola_ (fig. 33)

[Illustration: FIG. 32. _Hypocopra equorum_, perithecium with limited stroma, and detail of ascus tip with blue staining plug and spore. FIG. 33. Ascospores, from left, of _Sordaria fimicola_, _S. humana_ and _Sphaerodes fimicola_.]

114(3) Spores 20-28 × 12-16µm, with subacute ends, each with a germ pore. _Gelasinospora adjuncta_ - Spores larger. 115

115(114) Asci 4-spored. Spores 24-29 × 15-18µm, with rounded ends and one germ pore. _Gelasinospora tetrasperma_ - Asci 8-spored. Spores 26-3 _Gelasinospora cerealis_

116(4) Perithecia orange to golden, often gregarious, almost spherical, necks ca 50µm diam., 15µm high, setae at ostiole hyaline, up to 35 × 3µm. Spores limoniform, with a germ pore at each end, 15-25 × 9-16µm. _Sphaerodes fimicola_ (fig. 33) - Perithecia yellow or reddish brown (darker when filled with mature spores), neck 50µm long, with setae at the ostiole 40-70µm long. Spores dark brown to black, limoniform, 20-34 × 11-17µm, with apical germ pore. _Melanospora brevirostris_

117(2) Asci more than 8-spored. see Key 1 at 86 - Asci with 8 or fewer spores, or asci evanescent, not readily observed. 118

118(117) Perithecia orange/yellow, 500-1000µm diam. Spores long (over 45µm) or 2-celled if shorter. 119 - Perithecia smaller, or black or with a neck. Spores shorter (less than 20µm) or septate if longer. 120

119(118) Perithecia orange, 500-1000µm diam., in small groups on a limited stroma. Spores thick walled, 48-60 × 22-26µm, with a gelatinous sheath. _Selinia pulchra_ - Perithecia orange yellow, superficial, ca 500µm diam., with ostiole in a disc surrounded by silvery triangular tufts of hyphae ca 100µm long. Spores ellipsoid, 1-septate, 12-14 × 4-5µm. _Nectria suffulta_

120(118) Perithecia reddish brown or pale, hyaline, with a distinct neck. 121 - Perithecia black. 131

121(120) Perithecia globose, up to 250µm diam., immersed, reddish brown, with a neck 1-3 mm long. Asci broad ellipsoid, 5-8.5µm, rapidly breaking down and difficult to see. Spores ellipsoid-allantoid. 5.5-7 × 1.5-2µm, collecting in a pearly droplet at the fringed tip of the perithecial beak. _Viennotidia fimicola_ (fig. 34) - Perithecia pyriform, very pale in colour, 60-200µm diam., with a neck 60-700µm long. Asci rarely visible. Spores pointed-fusiform, 1-3 septate, often with a sheath and clumped together in fascicles. (_Pyxidiophora_, fig. 36) 122

122(121) Neck 95-145µm long, brown, rugose, with cells arranged in 5-6 longitudinal rows visible in one view. Spores 38-52µm long. _Pyxidiophora badiorostris_ - Neck not brown or rugose, composed of hyaline, irregularly arranged cylindrical cells. 123

123(122) Spores less than 45µm long. 124 - Spores more than 45µm long. 125

124(123) Spores 35-45µm long, with brown apical or subapical patches of pigment. _Pyxidiophora brunneocapitatus_ - Spores 35-43µm long, without brown apical or subapical patches of pigment. _Pyxidiophora microsporus_

125(123) Spores mostly 45-60µm long. 126 - Spores mostly longer than 60µm. 129

[Illustration: FIG. 34. _Viennotidia fimicola_, perithecium and spores. FIG. 35. _Phomatospora coprophila_, perithecium, and ascus with spores. FIG. 36. _Pyxidiophora petchii_, perithecium and spores.]

126(125) Perithecia 70-100µm diam., neck 100-190µm long. Spores (43)48-58(65)µm long. _Pyxidiophora grovei_ - Perithecia usually less than 80µm diam. 127

127(126) Perithecial necks mostly less than 100µm long. Spores (45)48-57(60)µm long. _Pyxidiophora arvernensis_ - Perithecial necks up to 200µm long. 128

128(127) Spores 45-53µm long. _Pyxidiophora petchii_ (fig. 36) - Spores 53-65µm long. _Pyxidiophora schotterianus_

129(125) Spores 60-70µm long. 130 - Spores (75)80-90(100)µm. Perithecia 120-160µm diam., neck 220-370µm long. _Pyxidiophora bainemensis_

130(129) Perithecial necks 300-700µm long. Spores 60-70µm. Perithecia 100-120µm diam. _Pyxidiophora spinuliformis_ - Perithecial necks 225-265µm long. Spores 65-70µm. Perithecia 110-125µm diam. _Pyxidiophora marchalii_

131(120) Perithecia small, up to 400µm diam., with hairy necks. Spores hyaline or pale, coppery-red en masse, extruded in tendrils. 132 - Perithecia larger, without hairy necks. If smaller than 200µm, with spores smaller than 5 × 3µm. 134

132(131) Spores reniform, with gelatinous sheath, 3.5 × 2-3µm, yellow, reddish brown _en masse_ in extruded tendrils. Asci spherical, evanescent. Perithecia black, spherical, 200-400µm diam., with cylindrical neck up to 300µm long, with sparse pointed hairs. _Microascus longirostris_ - Spores larger, not reniform. Perithecia up to 300µm diam. 133

133(132) Perithecial necks long, up to 750µm, with terminal hairs up to 1500µm, curved or circinate at tips. Spores limoniform, 7-10.5 × 5.5-7µm. _Lophotrichus ampullus_ - Perithecial necks short, ca 50µm, with long straight tapering hairs. Spore shape limoniform/variable, 6-7.5 × 5-5.5µm, with prominent germ pores. _Lophotrichus bartletti_

134(131) Perithecia up to 150µm diam., immersed but for a conical neck 50-75µm high. Asci 50 × 2-2.5µm. Spores minute, cylindrical, 3.5-4.5 × 1.75-2.5µm. _Phomatospora coprophila_ (fig. 35) - Perithecia more obvious, often hairy, or tomentose when young. Immature spores up to 70µm long, wavy cylindrical, with a row of globules inside and a short thin appendage at each end. (see _Cercophora_, 49)

135(1) Perithecia immersed, surrounded at the neck by a very limited flange-like stroma which is easily overlooked. see _Hypocopra_, 101 or if orange see _Selinia_, 119 - Stroma very conspicuous. 136

136(135) Perithecia in a subglobose group at the tip of the stromatic stalk. Spores with germ slit and gelatinous sheath. (_Podosordaria_) 137 - Perithecia not in a terminal head. 139

137(136) Stalk short, 3-5mm. Spores (12)14-19 × 6-9µm, slightly flattened on one side. _Podosordaria leporina_ - Stalk long, 1-6cm. Spores larger. 138

138(137) Spores 21-24 × 11-12µm. Stromatic stalk hairy. _Podosordaria tulasnei_ - Spores 40-60 × 20-30µm. Stromatic stalk not hairy. _Podosordaria pedunculata_

139(136) Stroma externally black, rooted or partially immersed in the dung, expanding at the surface to form a white disc up to 15mm diam., punctate with black perithecial ostioles. (_Poronia_) 140 - Stroma not as above. 141

140(139) Spores 18-26 × 7-12µm, bean shaped, with gelatinous sheath. Stroma deeply rooted. Especially on horse dung. _Poronia punctata_ - Spores (22)25-32(35) × (12)14-18µm, oblong ellipsoid to slightly fusiform. Stroma not deeply rooted. Especially on rabbit dung near the sea. _Poronia erici_

141(139) Stroma spreading over surface of dung or filamentous. Spores ellipsoid to slightly flattened on one side, with germ slit. (Xerophilic fungi developing after long periods of relatively dry incubation). (_Wawelia_) 142 - Stroma clavate, black, partly immersed to superficial, usually aggregated in small groups, ca 1-1.5mm high × 0.6-0.7mm diam., each containing a single perithecium. Spores ellipsoid with germ pore and gelatinous sheath. (_Bombardioidea_) 146

142(141) Stroma spreading on substrate, black brown, firm but not brittle. Ascomata globose, 0.5-1mm, with white hyphae at neck. Spores broad limoniform, 15-19 × 9-10µm. _Wawelia effusa_ - Perithecia globose to pyriform, black, brown or dark grey, produced laterally along the length of fine stromatal strand growing from the dung. 143

143(142) Asci 4-spored. 144 - Asci 8-spored. 145

144(143) Spores 15-18 × 9-12µm. Perithecia up to 400µm diam., dark grey at maturity, single or clustered, the ostiole with a crown of silvery white hyphae. Stromata up to 30 × 0.1-0.5mm. _Wawelia_ sp. - Spores 6-8 × 4-6µm. Stromata conical, white, 5-12 × 1-2mm. _Wawelia regia_

145(143) Perithecia hairy, globose, 350-500µm diam., stromatal strands up to 25mm long. Spores ellipsoid, flattened on one side, 9-12 × 6-8µm. _Wawelia octospora_ - Perithecia villose with conidiophores, globose, 230-420µm diam., produced laterally on stromatic filaments 20-30 × 0.1-0.3mm. Filaments pink at first, with a white pointed tip, becoming brown, velvety with conidiophores. Spores ellipsoid to flattened on one side, 7.5-9.5 × 3-4.5µm. _Wawelia_ sp. (fig. 37)

[Illustration: FIG. 37. _Wawelia_ sp., stromatic filaments with perithecia growing from a rabbit pellet, ascospores, and conidiophore and conidia.]

146(141) Asci 8-spored. Spores 20-31 × 9.5-15µm. _Bombardioidea bombardioides_ - Asci 4-spored. 147

147(146) Spores 24-34 × 15-19(20)µm. Basal germ pore less distinct than the apical one. _Bombardioidea serignanensis_ - Spores 34-43 × 16-22µm. Distinct germ pore at each end of spore. _Bombardioidea stercoris_

148 Fruit bodies solitary or in small groups, each a (key 1,1) subglobose, fertile, light brown head on a slender sterile stalk. Head soon bursting to expose the yellow ochraceous spore mass. On mixtures of bird droppings, cast pellets and decaying animal material. 149 - Fruit bodies superficial, lacking a distinct stalk. 150

149(148) Spores 5-8 × 2-3µm. Head 1-2mm diam. _Onygena corvina_ (fig. 38) - Spores 7-9 × 4-6µm, 4-5µm. Head 2-4mm diam. _Onygena equina_

150(148) Fruit bodies with an external wall of loosely anastomosing and interwoven hyphae, and with ± specialised terminal cells (GYMNOTHECIA, fig. 39). 151 - Fruit bodies with a well defined parenchymatic wall (CLEISTOTHECIA, fig. 46). 161

151(150) Gymnothecia with simple thin-walled, ± uniform and poorly developed hyphae constituting the outer hyphal sheath. 152 - Gymnothecia with thick-walled hyphae modified at their ends into appendages, or if thin-walled then always accompanied by appendages (i.e. curled, toothed or pointed hyphae). 155

[Illustration: FIG. 38. _Onygena corvina_, habit sketch, ascus and ascospore.]

152(151) Gymnothecia red-orange to brick-red. Ascospores orange, subglobose to ellipsoid, with an equatorial furrow, smooth, 4.5-5.5 × 3.5-4.5µm. _Arachniotus ruber_ (fig. 40) - Gymnothecia white or yellow, never orange or brick-red. Ascospores without an equatorial furrow. 153

153(152) Gymnothecia white. Ascospores hyaline, ellipsoid, smooth, 3-4 × 2-2.5µm. _Arachniotus candidus_ - Gymnothecia distinctly pigmented, yellow or brown. Ascospores larger than 4µm. 154

154(153) Gymnothecia yellow brown. Ascospores orange to brownish, slightly lenticular, smooth or slightly roughened, 5-6.5 × 3.3-4.6µm. _Arachniotus confluens_ - Gymnothecia lemon yellow. Ascospores lemon yellow, lenticular, smooth, 5-6 × 3-4.5µm. _Arachniotus citrinus_

[Illustration: FIG. 39. Habit sketch of a gymnothecium and ascus. FIGS 40-45. Spores and peridial hyphae. FIG. 40. _Arachniotus ruber._ FIG. 41. _Myxotrichum chartarum._ FIG. 42. _Gymnoascus californiensis._ FIG. 43. _Gymnoascus reesii._ FIG. 44. _Ctenomyces serratus._ FIG. 45. _Arthroderma curreyi._]

155(151) Gymnothecia possessing only thick pigmented hyphae. 156 - Gymnothecia possessing ± thin, hyaline hyphae with only a few, although often distinctive, appendages (i.e. comb-shaped end cells or dumb-bell shaped asperulate cells accompanying twisted and bent hyphae). 160

156(155) Gymnothecia brown-black or dark greenish-grey, with external hyphae with spine-like branches and septate, hooked appendages. Ascospores orange brownish, ovate, delicately striate, 4-5.2 × 2.4-3.3µm. _Myxotrichum chartarum_ (fig. 41) - Gymnothecia never black, and, if possessing thick-walled hyphae, then appendages never septate. Ascospores smooth, or if ornamented then asperulate or echinulate. 157

157(156) Gymnothecia rose to orange-brown or yellowish. Appendages curved or irregularly branched and pointed, never verticillately branched. Ascospores smooth, or at most asperulate. 158 - Gymnothecia red-brown with appendages verticillately branched. Ascospores 3-4.5 × 2-2.8µm, yellowish brown, lenticular. _Actinodendron verticillatum_

158(157) Gymnothecia rosy pink when young, becoming browner, with spines and curved, non-septate hairs. Ascospores hyaline, globose to subglobose, asperulate, 3-5 × 2.5-4µm. _Gymnoascus californiensis_ (fig. 42) - Gymnothecia yellow. Ascospores smooth. 159

159(158) Gymnothecia yellow to yellow-brown, without elongated appendages but with thick-walled branches, few of which are pointed. Ascospores globose-ellipsoid, yellow to brownish, 3-4.5 × 3.5µm. _Gymnoascus reesii_ (fig. 43) - Gymnothecia golden yellow to reddish-brown, with acute-ended appendages. Ascospores lenticular, smooth, hyaline, 2.5-3.5 × 2-2.5µm. _Pseudogymnoascus roseus_

160(155) Gymnothecia orange brown, with comb-like appendages. Ascospores slightly lenticular, pale orange, 3.3-3.6 × 2-2.6µm. _Ctenomyces serratus_ (fig. 44) - Gymnothecia whitish to pale ochraceous, particularly when dry, with few appendages but those present twisted and bent, and their branches constricted with regular or irregular dumb-bell shaped cells. Hyphal walls asperulate or with protuberances. Ascospores smooth, lenticular, hyaline, 2.4-3.3 × 2µm. _Arthroderma curreyi_ (fig. 45)

161(150) Asci relatively large, 100-200-spored, 1-3/fruit body. 'Cleistothecia' minute, <100 (rarely <250)µm diam., immersed. see _Thelebolus_ etc. (Key 1, 86) - Asci with 8 or fewer spores. 162

162(161) Ascospores purple at maturity, large, 50-70 × 25-35µm, epispore with a few longitudinal cracks. see _Ascobolus immersus_ (Key 1, 48) - Ascospores smaller, hyaline, yellow, olivaceous, brown or black. 163

163(162) Ascospores olivaceous, brown or black, at least in part. 164 - Ascospores aseptate, hyaline, yellow or other pale colours. 174

164(163) Ascospores 4-celled (cf. _Sporormiella_), with germ slits, readily fragmenting. Asci clavate, bitunicate. Cleistothecia black, shiny, up to 500µm diam. 165 - Ascospores 1- or 2-celled. 166

165(164) Ascus stalk up to 20µm long. Ascospores 25-32 × 5µm. _Preussia vulgaris_ - Ascus stalk 30-60µm long. Ascospores 26-38 × 5-7µm. _Preussia funiculata_ (fig. 47)

166(164) Ascospores 2-celled. 167 - Ascospores 1-celled. 170

167(166) Spores unequally 2-celled, one brown ellipsoid, with an apical germ pore, 10-12 × 6.5-7.5µm, the other a basal hyaline, cylindrical pedicel, 6-8 × 3µm. Cleistothecia black, globose, up to 250µm diam., covered with flexuous brown hairs up to 1mm long. Asci evanescent. _Zopfiella erostrata_ - Spores equally 2-celled. 168

168(167) Spores not constricted at the septum, ellipsoid, golden-brown, 25-30 × 10-15µm with 1-3 guttules in each cell. Cleistothecia gregarious on a mycelial mat, whitish to pale orange, up to 500µm diam. _Heleococcum aurantiacum_ (fig. 48) - Spores hyaline, divided into two almost globose cells by the constricting septum. Ascomata superficial, globose, dark coloured. (_Mycoarachis_) 169

169(168) Asci 8-spored, 5.5-11µm diam. Spores 5-5.5 × 3-3.5µm. _Mycoarachis inversa_ - Asci 4-spored, 6-6.5µm diam. Spores 4.5-5 × 2-2.5µm. _Mycoarachis tetraspora_

170(166) Asci broad-clavate, (1)-2-(3)-spored, 30-50 × 13-18µm. Spores brown-black with short ridges and warts, subglobose, 12-15.5 × 11-12.5µm, with a single germ pore. _Copromyces bisporus_ (fig. 49) - Asci 8-spored. 171

171(170) Spores globose, sooty brown, 3µm diam. Cleistothecia gregarious, with basal spirally coiled appendages, black, 100-200µm diam., partially immersed in a white to red felty hyphal mat. _Pleuroascus nicholsonii_ - Spores larger, ellipsoid or limoniform. 172

[Illustration: FIG. 46. Habit sketch of cleistothecia. FIGS 47-54. Asci and spores. FIG. 47. _Preussia funiculata._ FIG. 48. _Heleococcum aurantiacum._ FIG. 49. _Copromyces bisporus._ FIG. 50. _Arachnomyces nitidus._ FIG. 51. _Orbicula parietina._ FIG. 52. _Roumegueriella rufula._ FIG. 53. _Aphanoascus stercoraria._ FIG. 54. _Pseudeurotium ovale._]

172(171) Spores olivaceous, limoniform, usually with an apical germ pore. Perithecia greyish or greenish, abundantly hairy, branched or simple, straight or curly. Asci pedicellate, soon disappearing. see _Chaetomium_ at 5 - Spores darker, with 1 or more minute germ pores. Cleistothecia distinctly but not abundantly hairy. 173

173(172) Spores smoky brown, broadly ovoid, 9-14 × 6-9µm. Cleistothecial hairs short, up to 30µm. _Thielavia wareingii_ - Spores dark brown, flattened limoniform, 13-16 × 10-13 × 8-9µm. Cleistothecial hairs of two types, some smooth, dark brown, arising from the base up to 3mm long, others greyish green, rough, up to ca 120µm. _Thielavia fimeti_

174(163) Cleistothecia produced within a common arachnoid mycelial mass. Spores smooth or minutely asperulate, yellow to yellow-brown, broadly ellipsoid, 4-5 × 3-5µm. _Aphanoascus fulvescens_ - Cleistothecia single or gregarious, but not on or in a mycelial mass. 175

175(174) Cleistothecia 170-750µm diam., covered with long (several mm when extended), thick-walled, aseptate, helical appendages. Asci clavate cylindrical, evanescent, 35-62 × 12-21µm. Spores ellipsoid, hyaline, 12-17 × 9-12µm. _Lasiobolidium spirale_ - Cleistothecia without coiled appendages. 176

176(175) Cleistothecia with hairs or appendages. 177 - Cleistothecia smooth. 178

177(176) Cleistothecia black, shining, 100-200µm diam., with dark brown-black thick-walled hairs with hooked tips. Asci 8-15µm diam. Spores straw or copper coloured, ellipsoid, 4-7 × 3.5-4.5µm with de Bary bubble and a germ pore at each end. _Kernia nitida_ - Cleistothecia reddish brown, less than 1mm diam., with long simple appendages curled at the tips. Spores hyaline, oblate, 3.55 × 2-3µm. _Arachnomyces nitidus_ (fig. 50)

178(176) Ascospores globose, larger than 9µm. 179 - Ascospores ellipsoid, up to 9µm. Asci always subglobose. 180

179(178) Ascospores, smooth, 9-13µm. _Orbicula parietina_ (fig. 51) - Ascospores ornamented, 13-24µm Asci subglobose. Cleistothecia ochraceous, becoming yellowish brown or flushed cinnamon. _Roumegueriella rufula_ (fig. 52)

180(178) Ascospores hyaline, then faintly yellowish, minutely spiny, 2.5-3 × 2-2.5µm. Cleistothecia pale, then dark brown. _Aphanoascus stercoraria_ (fig. 53) - Ascospores hyaline, then brown, smooth, 5.5-6 × 3.5-4µm. Cleistothecia dark brown from the beginning. _Pseudeurotium ovale_ (fig. 54)

Key 3. Basidiomycota

1 Basidia single-celled (fig. 55). 2 - Basidia transversely or longitudinally septate (fig. 55), or difficult to observe. 71

2(1) Fruit body agaricoid, i.e. mushroom-shaped with gills underneath cap (figs 56, 67). 3 - Fruit body not agaricoid, without gills (figs 65, 66). 69

3(2) Spore print white or pale coloured, hyaline s.m. (Usually on straw/dung mixtures, never on raw dung except when very old). 5 - Spore print coloured. 4

4(3) Spore print pinkish or pale cinnamon, honey-coloured s.m. (Usually on straw/dung mixtures, never on raw dung). 6 - Spore print darker, in shades of brown or black. 8

5(3) Stem eccentric. Fruit body pure white. Spores ellipsoid, smooth. _Pleurotellus_ s. lato (If gills pink and spores longitudinally ridged see _Clitopilus passackerianus_, fig. 67) - Stem central. 7

[Illustration: FIG. 55. From left, sketches of holobasidium, with mature basidiospore showing germ pore; auriculariaceous basidium; tremellaceous basidium, lateral view and as often seen in sections.]

6(4) Fruit body white, ivory or very pale tan, with a smell of cucumber. Gills decurrent. _Clitocybe augeana_ - Fruit body yellow, with scaly cap. Gills free or just adnate. Fruit body with distinct ring and granular veil. (Commonly in plant pots. Probably associated with peaty material more than dung). _Leucocoprinus birnbaumii_ (_L. cepaestipes_ and _L. lilacinogranulosus_ occur in similar situations).

7(5) Fruit body with amethyst/purple shades, with eccentric stem. Spores subglobose, slightly ornamented to nearly smooth. (On compost heaps in gardens). _Lepista nuda_ - Fruit body with pink gills and distinct volva at stem base. Cap white to pale hazel. Stem white. Spores broadly ellipsoid, smooth. _Volvariella speciosa_

8(4) Spore print distinctly brown (fulvous, tawny, rust coloured etc.). 9 - Spore print some darker shade, fuscous, fuliginous or violaceous black. 20

9(8) Stem distinctly annulate, apex striate. _Conocybe percincta_ (Has been found on straw/dung mixtures, never on raw dung). - Stem lacking a veil. 10

10(9) Cap rich chrome yellow, viscid, soon reduced to a sticky mass, easily collapsing. _Bolbitius vitellinus_ - Cap in shades of brown, never brightly coloured and if collapsing then cap elongate-cylindric and white to pale cream. 11

11(10) Spore print dull, sepia or snuff-brown. On rabbit pellets in sand dunes. _Agrocybe subpediades_ - Spore print, brighter coloured, orange/rust brown. (_Conocybe_) 12

12(11) Gill edge with irregularly fusoid cystidia with obtuse apices (lageniform). Cap viscid. _Conocybe coprophila_ - Gill edge with distinctly capitate cells resembling a glass stoppered bottle (lecythiform). Cap never viscid, often pubescent under a lens. 13

13(12) Stem covered in long hairs. 14 - Stem covered in lecythiform cells similar to those on gill edge, giving a farinaceous appearance under a lens. NEVER with long hairs. (Dung/straw mixtures). Large as in a _Cortinarius_. Spores smooth. _Conocybe intrusa_ (_C. leucopus_ has been found on manured soil in gardens; _C. antipus_ has hexagonal spores and grows on dung piles).

14(13) Stem with both long hairs and lecythiform cystidia. [Illustration] 15 - Stem with hairs and lageniform cystidia. [Illustration] 16

15(14) Spores 11-14 × 7-9µm. Taste and smell strong, of fresh meal. _Conocybe farinacea_ - Spores large, over 15 × up to 10µm. Taste and smell none or slightly acidic. _Conocybe pubescens_ (_C. subpubescens_ might be found on straw/dung mixtures, and differs in spores 11-13 × 6-8µm).

16(14) Basidia 2-spored. _Conocybe rickenii_ - Basidia 4-spored. 17

17(16) Spores ellipsoid. 18 - Spores lentiform, angular in face view. _Conocybe lenticulospora_

18(17) Cap grey, contrasting with yellowish cream gills and pale stem. Spores 10.5-12.5 × 6-7µm. _Conocybe murinacea_ - Cap pinkish brown or tawny. 19

19(18) Spores 11-12 × 7.2-7.8µm. Cap sienna. On raw dung. _Conocybe fimetaria_ - Spores 10-12 × 6-7µm. Cap pinkish to cinnamon brown. On manured soil or sewage sludge. _Conocybe fuscomarginata_ (_Conocybe siennophylla_ might be found on straw/dung mixtures or in soil in greenhouses. It differs in having smaller spores).

20(8) Cap deliquescing to some degree at maturity. Basidia of 2 or 3 different sizes. (_Coprinus_) 21 - Cap not deliquescing. Basidia of one size only. 49

21(20) Veil on cap absent, cap either covered with small hairs (setules) or naked. 22 - Cap covered with a granular, micaceous, powdery or fibrillar veil. 28

[Illustration: FIG. 56. Habit sketch of a stipitate agaric, _Psathyrella stercoraria_, with section. FIG. 57. Sketch of gill section of _Psathyrella_ sp., showing position of marginal (m) and facial (f) cystidia. FIG. 58. _Coprinus pellucidus_, habit and vertical section of cap cuticle. FIG. 59. _C. pseudoradiatus_, habit and veil constituents. FIG. 60. _C. vermiculifer_, habit and veil constituents. FIG. 61. _C. filamentifer_, veil constituents. FIG. 62. _C. stercoreus_, habit. FIG. 63. _C. cordisporus_, vertical section of cap showing nature of veil cells on the cap cuticle. FIG. 64. Veil cells with structural (l) and superficial crystalline (r) ornamentation.]

22(21) Cap without setules. 23 - Cap with setules. 24

23(22) Cap minute, 1-5mm high before expanding, reddish orange at first, soon fading. Basidiospores almost globose to triangular in one view, elliptic in another, 7-10 × 7-9 × 5.5-6.5µm. (2- and 4-spored forms have been found). _Coprinus miser_ - Cap larger, up to 15mm when expanded. Basidiospores pip-shaped, 7.5-8.5 × 9.5-11 × 9.5-11.5µm. (4-spored). _Coprinus nudiceps_

24(22) Spores hexagonal, 10-13 × 6.5-7.5µm. Cap purplish. _Coprinus hexagonosporus_ - Spores ellipsoid. Cap brown or reddish, without purplish tints. 25

25(24) Basidia 4-spored. 26 - Basidia 2-spored. Spores 11-13 × 5.5-7µm. Facial cystidia absent. _Coprinus bisporus_ (_Coprinus sassii_, not yet recorded in British Isles, has 2-spored basidia with very large ellipsoid spores up to 20µm long).

26(25) Cap with a mixture of hyaline and brown thick-walled setules. Spores 9-10 × 5.5-6µm, with eccentric germ pore. Facial cystidia absent. _Coprinus heterosetulosus_ - Cap with only one type of setule. Facial cystidia present or absent. 27

27(26) Facial cystidia present. Spores 7.9-13.3 × 4.4-6.4µm, with apical germ pore. _Coprinus stellatus_ - Facial cystidia absent. Spores elongate and narrow, rarely greater than 5µm wide, with apical germ pore. Fruit body usually quite small, up to 6mm before expanding. _Coprinus pellucidus_ (fig. 58) (Several species in the group, e.g. _C. congregatus_ and _C. ephemerus_ have been found on straw/dung mixtures).

28(21) Veil strongly adhering to cap. Spores elliptic ovate, 15-20 × 8-12µm. Stem with distinct ring. Usually on buried dung. _Coprinus sterquilinus_ - Veil more floccose or powdery. Stem lacking ring or, if present (_C. ephemeroides_), fruit body small with 5-angled spores less than 10µm long. 29

29(28) Veil composed of filamentous units. 30 - Filamentous units, if present, masked by a preponderance of rounded cells. 35

30(29) Veil composed of strings of sausage-shaped, thin-walled, hyaline cells. 31 - Veil composed of rather narrow, slightly thickened hyphae. 32

31(30) Spores large, 11-14 × 6-7µm. Cap up to 1cm before expanding. Fruit body with or without a rooting base. _Coprinus radiatus_ - Spores smaller, up to 9µm long. Cap up to 6mm before expanding. Fruit body without a rooting base. _Coprinus pseudoradiatus_ (fig. 59) (_C. cinereus_ is found on straw/dung mixture and _C. macrocephalus_, with large spores, has been recorded on raw dung).

32(30) Veil citrus- or lime-yellow, or a mixture of hyaline and brown strongly coloured hyphae. 33 - Veil grey or whitish. 34

33(32) Veil of yellow hyphae. Spores 10.5-12.5 × 6-7.5µm. _Coprinus luteocephalus_ - Veil with brown hyphae. Spores 7-9 × 3.5-5µm. _Coprinus poliomallus_

34(32) Veil hyphae thin-walled. Spores 6.5-7.5 × 5µm, 'shouldered' about the apiculus. _Coprinus filamentifer_ (fig. 61) - Veil hyphae thin- and thick-walled, often with clamps. Spores elliptic-oblong, 9-10 × 5-6µm. _Coprinus vermiculifer_ (fig. 60) (_Coprinus flocculosus_, with spores 11.5-16.5 × 6-9.5µm, can be found on straw/dung mixtures).

35(29) Stem with small, distinct ring. Spores subglobose to lentiform and 5-angled, 6-9 × 6.5-8 × 5-6µm. _Coprinus ephemeroides_ - Stem at most with fibrils, even then rarely forming a faint ring zone. 36

36(35) with setules in addition to veil. 37 - Cap without setules. 38

37(36) Cap cystidia tapered. Spores 11-14 × 5-6.5µm. _Coprinus heptemerus_ - Cap cystidia capitate. Spores 10-11 × 6-7µm. _Coprinus curtus_

38(36) Veil of inflated bladder-like cells attached to filamentous units. Spores 7.5-8 × 4.5-5.5µm. _Coprinus utrifer_ - Veil of globose and subglobose cells and filamentous units often encrusted or with minute projections found sometimes at cap margin. 39

39(38) Globose cells, if ornamented then possessing crystalline or amorphous material (dissolved by 1N HCl, fig. 64.) 40 - Globose cells covered in small fine blunt projections on the walls (not removed by 1N HCl, fig. 64). 45

40(39) Basidia 2-spored. 41 - Basidia 4-spored. 42

41(40) Spores 14-17 × 8.5-10 × 12.5-14µm. _Coprinus pachyspermus_ - Spores smaller, 9-11 × 6-6.5 × 8-9µm. _Coprinus cordisporus_ (2-spored form)

42(40) Spores less than 10µm long. _Coprinus cordisporus_ (fig. 63) (_C. patouillardii_ is known on garden refuse, and an undescribed species with lemon-shaped spores has recently been found). - Spores 10µm or more long. 43

43(42) Veil soon discolouring greyish, drab or buff, Spores 11.5-14.5 × 6-8 × 7.5-9µm. _Coprinus cothurnatus_ - Veil remaining snowy white, only slowly discolouring greyish. 44

44(43) Fruit bodies several cm tall. Spores 15-19 × 8.5-11.5 × 11-13µm. _Coprinus niveus_ - Cap small, 5-6mm at first. Spores 14-16 x 8-9 × 10-12.5µm. _Coprinus latisporus_

45(39) Basidia 3-spored. 46 - Basidia 4-spored. 47

46(45) Spores narrow, 8.5-11 × 5-6.2µm. _Coprinus triplex_ - Spores broad, 9-10 x 6-6.5 × 6-7µm, slightly flattened in face view. _Coprinus trisporus_ (These are possibly a single taxon).

47(45) Spores 7-8 × 4-4.5µm, perispore not visible in water or alkali mounts. _Coprinus stercoreus_ (fig. 62) - Spores 9µm or more long. 48

48(47) Spores 9-11 × 5.5-6µm. Perisporal sac none or incomplete or indistinct. _Coprinus foetidellus_ - Spores longer, 10.8-13.5 × 5.5-7µm, with distinct perispore with dark lines and inclusions. Distinctive smell of gas. _Coprinus narcoticus_ (_C. sclerotiger_ is found on straw/dung mixtures, and the smaller _C. tuberosus_ on garden refuse etc.).

49(20) Spores not discoloured in conc. H2SO4. 50 - Spores discolouring in conc. H2SO4. Gills not spotted at maturity. 66

50(49) Cap cuticle cellular. Gills spotted at maturity. (More often on rich, 'dungy', soils. _P. subbalteatus_, with copper coloured cap, drying paler but retaining a dark marginal zone, occurs in gardens on mulch etc.). (_Panaeolus_) 51 - Cap cuticle filamentous. 56

51(50) Velar remnants very obvious, either as an appendiculate veil or as a distinct ring. 52 - Lacking all velar remnants. 54

52(51) Cap distinctly pigmented, with appendiculate veil. 53 - Cap pale coloured, smooth, semi-globate, soon cracking. Gills with marginal cystidia only. _Panaeolus papilionaceus_

53(52) Cap brown, smooth, sometimes viscid, not exceedingly wrinkled. _Panaeolus campanulatus_ - Cap grey, olivaceous, even black, with contrasting white appendiculate veil. _Panaeolus sphinctrinus_

54(51) Cap with or without appendiculate veil, but always with distinct ring. _Panaeolus semiovatus_ - Cap lacking veil. 55

55(54) Cap pinkish ochraceous to tawny-buff. Lacking facial cystidia. _Panaeolus speciosus_ - Cap whitish or slightly yellowish. With facial cystidia. _Panaeolus antillarum_

56(50) Gills with facial cystidia often containing yellow amorphous material when seen in ammonia solution or deep blue with cotton blue. (_Stropharia_) 57 (Blue-green _S. cyanea_ & _S. aeruginosa_ often occur in rich garden soils). - Gills lacking facial cystidia. Never with yellowing cystidia in ammonia. (_Psilocybe_) 58 (Red-capped _P. aurantia_ can be found on straw/mulch mixtures in gardens).

57(56) Cap sticky, semi-globate ± expanding at maturity. On raw dung. _Stropharia semiglobata_ - Cap plano-convex, often broad with a central umbo, margin flaring with age. On dungy mixtures in gardens. _Stropharia stercoraria_

58(56) Stipe bluing, with ring. Spores ellipsoid, 11-14 × 6.5-7.5µm. Fruit body with mealy smell and taste. _Psilocybe fimetaria_ - Stipe lacking distinct ring, or if with ring or ring zone 2-spored and/or stem not bluing. Fruit body without mealy smell and taste. 59

59(58) Stem always with distinct ring. Basidia 2-spored. Spores 15-20µm long. _Psilocybe luteonitens_ - Stem with or without ring. Basidia 4-spored. If with ring, spores smaller. 60

60(59) With ring zone. 61 - Lacking velar remnants on stem, or only appendiculate teeth at cap margin. 62

61(60) Spores slightly angular/limoniform, 11-13(14) × 7-8µm. Often on sewage sludge. _Psilocybe merdaria_ - Spores 13-14 × 7.5-8.5µm. _Psilocybe moelleri_

62(60) Spores 14-20 × 8-10µm. _Psilocybe subcoprophila_ - Spores smaller. 63

63(62) Spores lentiform, angled, 6-8(8.5) × 4.5-5.5 × 3.75-4.5µm. _Psilocybe bullacea_ (_P. crobula_, occasional on dung, differs in lacking purple colour in gills, and slightly smaller, ovoid, not angular, spores). - Spores larger. 64

64(63) Spores ellipsoid to slightly amygdaliform. _Psilocybe merdicola_ - Spores lentiform, angular. 65

65(64) Spores 11-13(14) × 7-8(9)µm. see _Psilocybe merdaria_, 61 - Spores 12-15 × 8-9.5µm _Psilocybe coprophila_

66(49) Round cells on cap as a micaceous veil. (Re-examine gill face; if different sized basidia and facial cystidia separating the gills are present go to _Coprinus_ at 21). _Psathyrella sphaerocystis_ - Cap lacking veil, or if present then fibrillar. 67

67(66) White copious veil at margin or also covering cap centre. Spores 10-12 × 5.5-6µm. _Psathyrella coprobia_ - Lacking copious veil. 68

68(67) With red edge to gill. Spores 12-13 × 6-6.5µm, with central germ pore. _Psathyrella stercoraria_ - Lacking red gill edge. Spores with eccentric germ pore. _Psathyrella coprophila_ (_P. fimetaria_ differs in spore size; there are several members of the _P. prona_ group which grow on soil/straw mixtures).

69(2) Fruit body club-shaped. _Typhula setipes_ (fig. 65) (_Clavaria acuta_ often grows on peaty soil in pots in greenhouses). - Fruit bodies effuse, resupinate 70

70(69) Fruit-body cobweb-like and greyish white. Basal hyphae 3-4.5µm wide. Spores sub-globose, 4.5µm diam. (Generally on old dung or straw/soil mixtures). _Athelia coprophila_ (If with spiny spores 5-6µm diam., see the recently recorded _Tomentellopsis echinospora_). - Fruit-body with pores, white or flushed slightly ochraceous, brownish or greyish. (On clods of soil in dunged land). _Cristella candidissima_

71(1) Fruit body either a cup containing several 'eggs' or a single orange or yellowish gelatinous sphere. 72 - Fruit-body effuse, without distinct shape. 73

72(71) Fruit-body whitish or pale yellow, up to 2.5mm diam., splitting at maturity to shoot away the orange/yellow spore mass. _Sphaerobolus stellatus_ (fig. 66) - Fruit-body cup shaped, with silvery-grey 'eggs'. (Usually on dung and straw or attached to rabbit pellets). _Cyathus stercoreus_ (_Cyathus vernicosus_ often grows in plant pots on rich soil).

73(71) Basidia with transverse septa. Spores 11 × 7µm. Fruit body pinkish. _Platygloea fimicola_ (Not British; included for completeness. _Pilacrella solani_, with a glistening stipitate head, has been isolated from dungy soil). - Basidia with longitudinal septa. Spores 14-18 × 9-10µm. Fruit body cream-white or ivory. _Sebacina incrustans_

[Illustration: FIG. 65. Habit sketch of _Typhula_ sp. Note attachment to sclerotium. FIG. 66. _Sphaerobolus stellatus_, habit. FIG. 67. _Clitopilus passackerianus_, a sessile agaric--habit sketch and section.]

Key 4. Zygomycota

1 Spores formed in multispored sporangia (figs 68, 70, 72, 75, 76) or in few-spored sporangioles (figs 70, 73). 2 - Multispored sporangia and globose sporangioles absent. Spores formed singly on terminal, lateral or intermediate vesicles (figs 74, 79, 80, 82-86), or in short chains (figs 77, 78, 81). 11

2(1) Sporangiophore stout, simple, with a subsporangial swelling and a basal swelling buried in the substrate. Sporangia tough walled, black, projected some distance towards the light when mature, and sticking to whatever they hit. _Pilobolus_ (fig. 76) e.g. spores pale yellow, 8-10 × 5-6µm - _P. crystallinus_ spores orange, 12-20 × 6-10µm. - _P. kleinii_ - Sporangiophores not stout; sporangia not violently discharged. 3

3(2) Sporangial wall black, tough, not readily broken when touched. Sporangia with a sticky base, becoming attached to whatever they contact after the marked elongation of the white sporangiophores at maturity. _Pilaira_ (fig. 75) e.g. spores yellowish, 8-10 × 6µm - _P. anomala_ spores colourless, 11-13 × 6-8µm - _P. moreaui_ - Sporangial wall diffluent, spores readily removed in a droplet, or fragile and then spores easily dispersed by external violence. 4

4(3) Sporangiophores stiff and metallic in appearance, growing towards the light and often to great length (5-30cm). _Phycomyces_ e.g. spores 10.5-30 × 6.5-17µm; columella pyriform; sporangiophores up to 30cm - _P. nitens_ spores 8-13 × 5-7.5µm; columella spherical or ovoid; sporangiophores up to 30cm - _P. blakesleeanus_ - Sporangiophores white, not reaching extreme lengths. 5

5(4) Small lateral sporangia (sporangioles) present. 10 - Sporangioles absent. 6

6(5) Sporangiophores usually grouped, less often single, connected by stolon-like hyphae. 7 - Sporangiophores arising singly, or if grouped then lacking stolon-like hyphae. 9

7(6) Stolons joining groups of sporangiophores often with rhizoids at the base of the group. 8 - Sporangiophores arising singly or in groups from stolons, which may be 'rooted' at intervals along their length, but rarely beneath the groups of sporangiophores. _Absidia_ (fig. 71) e.g. sporangiophores grouped, rhizoids poorly developed; spores 2.5-4.5µm diam. - _A. corymbifera_ sporangiophores grouped, rhizoids strongly developed; spores 2.5-3.5µm diam. - _A. orchidis_

8(7) Sporangiophores mostly unbranched. _Rhizopus_ (fig. 69) e.g. spores irregularly angular-ovoid, 8-14 × 11µm - _R. nigricans_ - Sporangiophores with a whorl of branches beneath the main sporangium, each with a small columellate sporangium. Spores 6-8.5µm. _Actinomucor elegans_

[Illustration: FIG. 68. _Mucor_, habit and detail of sporangium before and after dehiscence. FIG. 69. _Rhizopus_, habit. FIG. 70. _Thamnidium elegans_, habit and detail of sporangioles. FIG. 71. _Absidia_, habit. FIG. 72. _Mortierella_, habit and sporangiophore tip after sporangial dehiscence. FIG. 73. _Helicostylum_, sporangioles. FIG. 74. _Chaetocladium_, sporangioles. FIG. 75. _Pilaira_, sporangiophores before and after elongation, and sporangium. FIG. 76. _Pilobolus_, sporangiophore. FIG. 77. _Syncephalis_, habit, sporangiophore and merosporangia. FIG. 78. _Piptocephalis_, habit and detail of final branch with head cell and merosporangia. FIG. 79. _Oedocephalum_, habit and sporing head. FIG. 80. _Rhopalomyces_, sporing head. FIG. 81. _Syncephalastrum_, habit and detail of merosporangium. FIG. 82. _Coemansia_, habit, sporoclade with sporangia and sporangium with spore inside. FIG. 83. _Kickxella_, habit and sporoclade. FIG. 84. _Cunninghamella_, habit and fertile head. FIG. 85. _Mycotypha_ (l) and _Ostracoderma_ (r) conidiophores. FIG. 86. _Ballocephala_, habit of sporangiophores growing from parasitised tardigrade, sporangiophore and sporangia.]

9(6) Sporangia often with pigmented walls, yellowish when young, finally grey or black, with well marked columella left after spore dispersal. Individual sporangiophores observable with unaided eye, up to 20mm long. _Mucor_ (fig. 68) e.g. spores smooth, 7-8 × 2.5-4.5µm - _M. hiemalis_ spores smooth, 6-12 x 3-6µm - _M. mucedo_ spores asperulate, 5-8.µm diam. - _M. plumbeus_ (N.B. _Zygorhynchus_ would key out with _Mucor_. It is more often isolated from soil, and is distinguished from _Mucor_ by the presence of zygospores with unequal suspensors) - Sporangia white, without a columella, readily becoming a spore droplet. Sporangiophores delicate, often only 200-400µm long. Fine, white, garlic-smelling mycelium often present. _Mortierella_ (fig. 72) e.g. spores 16-27µm diam, few in each sporangium; sporangiophores ca 150µm, with short lateral branches at right angles - _M. reticulata_ spores 6-10 x 4-6µm; sporangiophores 2-3mm high, with ascending branches - _M. bainieri_ spores 4-10µm; sporangiophores richly branched - _M. candelabrum_

10(5) Sporangioles formed at the final tips of a densely dichotomous system of branchlets, originating some distance below a terminal sporangium (which may be absent in young specimens). Sporangioles up to 25µm diam., with up to 6 spores. Spores 8-12 × 6-8µm. _Thamnidium elegans_ (fig. 70) - Sporangioles either at the curved tips of slender branches, or clustered in groups about halfway along tapering branches which radiate from the sporangiophore below the sporangium; the branch tips of the latter give the fertile portion of the sporangiophore a bristly appearance. _Helicostylum_ (fig. 73) e.g. spores 8-17 × 3-7µm; sporangioles on short secondary or tertiary branches; fertile region bristly with sterile branches - _H. fresenii_ spores 6-8 × 4µm; sporangioles reflexed, on slender primary or secondary branches; fertile region without sterile branches - _H. pyriforme_

11(1) Spores formed in chains. 12 - Spores formed singly. 14

12(11) Sporangiophores regularly and repeatedly dichotomously branched. Chains of 2-10 spores produced in small groups, which may be wet or dry, on deciduous heads, 4-15µm diam. Parasitic on other fungi, mostly other Mucorales. _Piptocephalis_ (fig. 78) e.g. spores 4-5 × 2-3µm, in pairs; heads dry - _P. lepidula_ spores 5-6 × 2-2.5µm, in chains of 4-9; heads dry - _P. cylindrospora_ spores 4-8 × 2-4µm, in chains of 3-5; heads dry; sporangiophore without rhizoids - _P. freseniana_ spores 4-6 × 4-4.5µm, in chains of 3-6; heads wet; sporangiophore with rhizoids - _P. repens_ spores 3-5 × 2-2.5µm, in chains of 3-5, heads wet; head cell lyses, to leave only a fringe at the tip of the very fine sporangiophore - _P. fimbriata_ - Sporangiophores simple or irregularly branched. 13

13(12) A large conspicuous fungus, macroscopically Mucor-like, mycelium coarse. Sporangiophores with a distinct terminal swelling with crowded spore chains. Spores usually 5-10 in a chain, globose to ovoid, 2-8 × 4-6µm. _Syncephalastrum racemosum_ (fig. 81) - Sporangiophores less conspicuous, 100-1000µm high, with a 'holdfast' at the base attaching the sporangiophore to the substrate. Mycelium very fine. Parasitic on other Mucorales. _Syncephalis_ (fig. 77) e.g. sporangiophores 100-200µm high, with three 'nodes' along their length; merosporangia often forked at the basal cell; spores 8-10 × 6µm - _S. nodosa_ sporangiophores up to 750µm high; merosporangia usually subdivided at their base into several branches, each with 5-10 spores; spores 5-10 × 3-4µm - _S. depressa_ (N.B. _Oedocephalum_ spp. (fig. 79), the anamorphic states of many dung fungi (esp. Ascobolaceae and Pezizaceae), _Rhopalomyces_ (fig. 80), and some _Aspergillus_ spp. are superficially similar to _Syncephalis_ at first sight).

14(11) Sporangia containing a single closely fitting elongated spore, produced in serried ranks on one side of a boat-shaped branch (sporoclade). 15 - Single-spored sporangia ('spores') globose, produced singly or if in groups not on sporoclades. 16

15(14) Sporoclades lateral. Sporangiophores usually yellowish. (No parasitism has been demonstrated, but in culture grows much better in the presence of the white, garlic-smelling _Mortierella_ spp.). _Coemansia_ (fig. 82) e.g. spores 6-11µm long; sporoclades spirally arranged around the axis - _C. erecta_ spores 16-18µm long; sporoclades formed on one side of the axis, causing it to curve to one side - _C. scorpoidea_ - Sporoclades produced in a terminal verticil. Sporangiophores shining white. _Kickxella alabastrina_ (fig. 83)

16(14) Spores produced in clusters below the apex of the final branches of a compound, often trifid, branching system which is given a bristly appearance by the projecting tips. Superficially similar to _Thamnidium_ or _Helicostylum_. Capable of parasitising, and growing much better in association with, other Mucorales. _Chaetocladium_ (fig. 74) e.g. spores smooth, 4-6µm diam. - _C. brefeldii_ spores echinulate, 6.5-9.5µm - _C. jonesii_ - 'Spores' not produced in subterminal clusters, but terminally on lateral vesicles, or over the surface of swollen fertile regions of the sporangiophore. 17

17(16) Sporangiophores up to 250µm high. Lateral vesicles numerous, each producing a single 'spore', which is projected when mature. Parasitic on tardigiades. _Ballocephala_ (fig. 86) - Sporangiophores visible with the unaided eye. Spores produced on swollen parts of the sporangiophore. 18

18(17) Sporangiophores branched, with more or less globose terminal fertile regions. Spores dry and powdery, yellowish or pinkish in mass. _Cunninghamella_ (fig. 84) e.g. spores smooth, ovoid, 18-22 × 10-14µm or globose, 8-10µm diam. - _C. elegans_ spores echinulate, ovoid, 8-12µm - _C. africana_ - Sporangiophores unbranched, fertile portion 200-300 × 15-20µm. Fertile region terminal only, cylindrical. Spores smooth, greyish in mass, 2-4µm diam. _Mycotypha microspora_ (fig. 85) (N.B. _Ostracoderma epigea_ (fig. 85), the anamorph of _Peziza astracoderma_, which occurs on paper and sometimes dung and highly organic substrates, was originally described as _Mycotypha dichotoma_. The fertile regions are cylindrical but multiple as the result of several close dichotomous divisions at the base of the fertile portion).

Notes

[1] There are few reports of _Ascozonus_, apart from _A. woolhopensis_. Observed spore sizes of _A. woolhopensis_ suggest that measurement of Renny's (1874) illustrations of spores leads to values which are too large (19-20 × 6-6.5µm). Those in parentheses are what they might be, based on the discrepancy between observed values for _A. woolhopensis_ and Renny's illustration.