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A key to reddening species of Inocybe in Britain
E L F GIH
A key to reddening species of Inocybe in Britain >LL @
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hen working through a dichotomous key to a genus having a large number of species such as Inocybe, it can be somewhat frustrating to have to negotiate a long and often tortuous journey when one’s specimen has a particularly striking character. It can be tantalising to find no reference to this character along the way until perhaps just a brief mention at journey’s end. Mind you, having dabbled to some extent with devising such keys I do have sympathies for the challenged designer as well as the sometimes sorely tested user. It was with this issue in mind that I decided to design a key to the species of Inocybe that redden when bruised.
Subgenus Inosperma Section Rimosae (to which these two species belong) probably contain some degree of muscarin. I. erubescens, another reddening species of subgenus Inosperma, has sufficient quantities of muscarin to render it deadly poisonous. Bearing in mind the considerable danger of consuming I. erubescens in error, it makes good sense for those of us who enjoy collecting for the pot never to eat (or even taste) any member of this genus. In any case, taste is not amongst the characters used in identification in Inocybe. Microscopy It is always essential to observe microscopic characters when working with this genus, and I make no apology for referring to spore shape and cystidia characteristics right from the start of the key. If this is not your forte then read no further – making reliable determinations of Inocybe without recourse to a microscope is unfortunately not possible. For examination the only chemical needed is 10% ammonia, no other stains are necessary, and hymenial cystidia (those found on the gill) are quite easy to locate compared to some other genera. Another plus: the fruitbodies as a rule are quite resilient and will normally survive for several days in the fridge or even in the post, the only proviso being that as with most agarics a sporeprint for really accurate spore measurement needs to be acquired from an unrefrigerated specimen. For the purposes of this key, only cheilocystidia (found on the gill edge) need be examined: if these are thin-walled one can safely assume that pleurocystidia (found on the gill face) are absent; conversely if thick-walled and capped with crystals (metuloids) one can safely assume that pleurocystidia are also present (if both sorts are absent on the gill edge then one probably has a different genus).
Types of reddening Of the approximately 140 species of Inocybe in Britain, 15 show reddening in some part of the fruitbody to varying degrees. The reddening itself varies from species to species and can be bright red, vinaceous, orange-red to rust-red: it can occur in some species throughout the fruitbody, in others just in the stipe. In two species, I. corydalina and I. erinaceomorpha, reddening is only rarely present in my experience, but they are included here for completeness. In all but two, I. adaequata and I. rhodiola, where the dark vinaceous red staining found in the cap, stipe and flesh seems to be inherent from the start, the reddening is not present in young, pristine specimens but develops either as they mature or with handling or damage. Toxins Though many species of Inocybe (including some of the reddening species) are reported to contain dangerously poisonous toxins, including psilocybin and muscarin, Kuyper (1986) comments that one of the reddening species, I. adaequata [therefore possibly I. rhodiola also?], contains no muscarin and is considered the only safely edible member of the genus. All other members of
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doi:10.1016/j.fldmyc.2015.06.004
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E L F GIH Caulocystidia Stipe pruinosity and locating caulocystidia correctly is generally much harder to assess and (in my experience) is the commonest cause of errors and confusion in identification of Inocybe, though only the second half of this key requires this information. I have found that even with the best dissecting scope it is not always possible to judge safely whether pruinosity apparent on the stipe is due to true caulocystidia or merely hairs (surface hyphae which become detached to some degree). The only safe way is to take the finest strip of fibre from the stipe and examine it under a compound microscope in ammonia, at the same time noting from exactly which part of the stipe your strip came. To use the key you may need to know whether caulocystidia are: 1) absent even at the apex, 2) present only at the apex, or 3) present in the top 3rd or in one species right down to the base. A word of caution: it is all too easy to remove or damage caulocystidia through overhandling on collection, thus giving you a false reading when assessing with a microscope. An added benefit to making up a slide from the stipe is that the spores found there will have dropped naturally and will therefore be mature. Thus in the absence of a spore print this gives you more accurate measurements than if measured from a gill squash.
included therein, also that I. rhodiola though included is not there accepted as British. This is rectified in Edition 3 (Outen & Cullington, 2015), now available online. Both species are recognised by some (if not all) authorities in Europe and have now been found in Britain. At present with this genus we are reliant upon workers abroad whose research using DNA is proving extremely useful. Until such time as British collections of Inocybe can be similarly researched there will inevitably remain an element of doubt over declaring new species here. Species descriptions A fairly detailed description follows each species as it occurs in the key. Please note, however, that to avoid unnecessary repetition the main characters already covered in the relevant couplet are not then duplicated in the description unless further explanation is useful. Stature (medium, large etc) is relative to members of the genus as a whole. Inocybe Key For complete British coverage of this genus there is now a new edition of Outen and Cullington’s Keys to British species of Inocybe (Outen & Cullington, 2015), available free of charge online, hosted by the Bucks Fungus Group’s website: (www.bucksfungusgroup.org.uk). Edition 3:1 April 2015 (Cullington) is much changed from previous editions with, amongst other things, a complete revision of Key D for nodulose-spored marginate species, also the key to reddening species included in this article. This latest version is recommended in place of previous editions. To gain access you will need to email me for the necessary code: ([email protected]).
Measurements of cystidia and spores With this genus the size and shape of cystidia (and to a lesser extent spores as well) can vary far more than is met with in some other genera even within one gill, let alone one specimen. Re size: I find the best approach is to make note of the longest / widest / thickest-walled examples rather than attempt to assess an average, then compare this with the greatest measurement given in a description (bearing in mind that cystidia found on the gill face tend to be slightly bigger than those on the edge). The same advice applies to measuring spores. Regarding cystidia shape: it is well-nigh impossible to cover in a nutshell the variety of shapes which may occur in one gill preparation; I advise aiming for an overall general impression, taking more note of the larger examples which tend to be more typical.
All photographs of fruitbodies and microscopy are © Penny Cullington unless otherwise attributed.
Two new species Owners of the printed versions of Outen & Cullington Keys will note that I. armeniaca is not
/3
E L F GIH .+0 -0. . , 0 .(,. 1
Spores nodulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. bresadolae Fruitbody of medium stature; cap to 45(50) mm, campanulate then flat with an umbo, tomentose to ± scalyfibrillose, pale ochraceous brown with orange tint, reddening near margin; gills beige, eventually reddening; stipe to 45 × 7 mm, yellowish, sometimes reddening, sometimes with (marginate) bulb, entirely pruinose with caulocystidia; flesh with guaiac stains intense wine-red; smell sweetish as in I. bongardii; spores 7–10 × 5–7 µm with a few small nodules; hymenial cystidia metuloid, fusiform, to 75 × 17 µm, walls to 3 µm thick, yellow in ammonia; under deciduous trees in summer, reportedly rarely in autumn. Rare: very few British records. (Figs 27, 28).
1*
Spores smooth – either ellipsoid to phaseoliform or amygdaliform . . . . . . . . . . . . . . . . . . . . . . . . 2
2 2*
Spores ellipsoid to phaseoliform; cheilocystidia thin-walled, pleurocystidia absent . . . . . . . . . . . 3 Spores amygdaliform; cheilocystidia thick-walled, mostly crystal-tipped, pleurocystidia present 8
3
Cap predominantly scaly, squamulose to squarrulose, some shade of brown; rust-red staining confined to stipe where handled or damaged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Cap smooth to markedly fibrillose, never becoming predominantly scaly, either whitish or vinaceous brown; reddening occurring throughout fruitbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3*
4
Cap scales darkish brown on a paler brown ground, becoming ± squarrulose at centre; smell strong and disagreeable, earthy or musty; mainly under Picea . . . . . . . . . . . . . . . . . I. cervicolor
Fruitbody of medium stature; cap to 45 mm, ± domed then flat; gills pale yellowish then brown; stipe to 100 x 6 mm, concolorous, ± flocculose above, streaky fibrillose below where becoming rust-red when handled; spores smooth, ellipsoid to subphaseoliform, 11–15 × 6.5–8.5 µm; cheilocystidia thin-walled, wavycylindrical to slenderly clavate, to 50(60) × 14(16) µm, pleurocystidia absent. Uncommon. (Figs 25, 26).
4*
Cap scales mid-brown on a paler buff to pinkish brown ground, remaining ± squamulose; smell either pleasant and fruity, or if unpleasant then fishy; mainly under Fagus and Quercus . . . . . . 5
5
Smell sweetish with flowery or fruity component. . . . . . . . . . . . . . . I. bongardii var. bongardii
Fruitbody of medium stature though stipe sometimes disproportionately long; cap to 50(60) mm, ± campanulate; gills unremarkable greyish brown; stipe to 80(90) × 9 mm, concolorous, ± flocculose above, streaky fibrillose below where becoming rust-red when handled; smell usually pleasant (though when found with Helianthemum smell faint, ± earthy?); spores smooth, ellipsoid to subphaseoliform, 11–16 × 6.5–9 µm; cheilocystidia thin-walled, wavy-cylindrical to slenderly clavate, to 60 × 18 µm, pleurocystidia absent; under Fagus, Quercus and Castanea, also with Helianthemum; occasional. (Figs 22, 24).
5*
Smell unpleasant, slightly fishy cf. Macrocystidia cucumis . . . . . . I. bongardii var. pisciodora Details as for the type variety above, but smell complex: reportedly sometimes with a Pelargonium component and at some stage becoming unpleasant and fishy. It is advisable to contain specimens and check periodically after collection to assess the smell which can be elusive, even ephemeral. Rarely reported; I have found this variety under Fagus.
6
Cap smooth; cap and stipe initially white, soon becoming ochraceous to flesh-coloured, developing bright red stains with age and/or damage; smell ± sickly, of soap; fruiting in June or July under Fagus on calcareous soil. Dangerously poisonous. . . . . . . . . . . . . . . . . . . . . . . . . . . . I. erubescens Fruitbody of medium to large stature; cap to 70 mm or more, at first with abruptly downturned margin, then with obtuse umbo, splitting with age; gills pale grey-beige then dirty olivaceous, bruising red where touched; stipe to 70(90) × 15(20) mm, concolorous, sometimes with clavate base; flesh off-white to pinkish, bruising red; spores smooth, ellipsoid to subphaseoliform, to 14 × 7.5 µm; cheilocystidia slightly clavate, to 65 × 15(20) µm, pleurocystidia absent; uncommon (Figs 21, 23).
6*
Cap markedly streaky fibrillose, cap and lower stipe dirty russet to vinaceous brown, upper stipe remaining paler; flesh discolouring pinkish to vinaceous from the start; smell different; fruiting in autumn under various deciduous trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
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E L F GIH 7
Fruitbody robust, cap to 60(80)mm with obtuse umbo and stipe to 70 x 12 µm; maturing dirty brownish purple; smell unpleasant, earthy or of beetroot; not uncommon especially under Quercus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. adaequata
Cap streaky fibrillose, sometimes furrowed, dirty vinaceous red-brown; gills at first pale beige then darker with reddish tinge; stipe ± cylindrical, not pruinose, not bulbous, pale at first ( almost white at apex), then from the base increasingly pinkish to dark reddish brown; stipe and cap flesh white then soon pinkish, becoming darker vinaceous brown from stipe base upwards; spores smooth, ellipsoid to slightly phaseoliform, 11–13(14) × (5)6–7(8) µm; cheilocystidia thin-walled, wavy-cylindrical, to 70 × 15 µm; under deciduous trees, usually Quercus; occasional. Delimitation between I. adaequata and the smaller but similar I. rhodiola (shown to be distinct with DNA, Larsson 2009) is not yet clear, although I. rhodiola appears to retain its pink stipe tones rather than darkening to vinaceous brown; several British collections matching well to I. rhodiola have been made since then. (Figs 17, 18).
7*
Fruitbody smaller, more delicate and slender, umbo acute to obtuse; stipe usually pinkish below; smell faint; probably rare, found so far under Fagus in S. England . . . . . . . . . . . . . . . I. rhodiola Microscopic details are as far as known similar to I. adaequata. See comments re delimitation under that species. (Figs 19, 20).
8
Stipe entirely pruinose with caulocystidia, base marginately bulbous; cap pale yellowish but fruitbody becoming increasingly stained bright red, sometimes ± entirely so when old . . I. godeyi Fruitbody of small to medium stature; cap to 40(50) mm, conical then with ± acute umbo, finely fibrillose, sometimes scaly with age, almost white then ochraceous but soon strongly (even entirely) reddening; gills pale beige then ochraceous, reddening; stipe to 40(50) × 6 mm, concolorous, reddening; flesh pale but reddening; smell acidic; spores smooth, amygdaliform, to 12 × 7 µm; hymenial cystidia metuloid, swollen fusiform, to 65(70) × 25 µm, walls to 1.5(2) µm thick; under deciduous trees on calcareous soil; occasional, tending to fruit in summer to early autumn. (Figs 11, 13).
8*
Stipe either pruinose with caulocystidia descending no lower than the top third, or caulocystidia virtually absent though may appear pruinose/hairy at apex, base not bulbous . . . . . . . . . . . . . . . 9
9.
Cap white and ± silky smooth, developing orange-red staining as it matures, especially where handled; stipe white with tendency to redden, pruinose with caulocystidia in top 3rd (sometimes less far down?); smell unremarkable, spermatic; spores with ± obtuse apex; under conifers and deciduous trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Cap some other colour and not silky smooth, either fibrillose or scaly; stipe appearing pruinose at apex but caulocystidia virtually absent; smell unusual: often strong, fruity, of pear-drops or of urine/stables; spores with ± conical apex; only under deciduous trees . . . . . . . . . . . . . . . . . . . . . 11
9*
10
Fruitbody small and slender, similar in appearance to I. geophylla but additionally with orangered staining developing on part of cap and / or stipe. Rare. . . . . . . . . . . . . . . . . . . . . I. armeniaca
Cap to 20(30) mm, silky smooth to finely fibrillose, conical usually with acute umbo when young, white to palest cream but developing orange-red patches which separate it from I. geophylla in the field; gills unremarkable, possibly also orange-red where damaged; stipe to 50 × 2(5) mm, ± cylindrical, pruinose near apex, cortina present when immature, not bulbous, concolorous, possibly becoming orange-red with age; flesh whitish, also possibly becoming orange red with age; smell if present spermatic; spores smooth, amygdaliform with rounded apex, av. 9 × 5 µm; hymenial cystidia metuloid, similar to those of I. geophylla, walls to 2 µm thick, pale yellow in ammonia; misshapen caulocystidia present near stipe apex amongst cauloparacystidia which extend further down (though according to Bon caulocystidia absent?). Specific tree association in Britain is as yet not established; in Europe reported from both sandy and calcareous soil under a mix of conifers and deciduous trees. I have seen two collections in Britain (see Field Mycology Vol 15(4): 134) though sadly the material was not retained. Further collections are needed to clarify species details and provide fungarium specimens to back up its presence here. (Figs 15, 16).
32
E L F GIH 10*
Fruitbody larger, of medium stature, cap to 40 mm or more, ± domed, rarely with acute umbo, i.e. not reminiscent of I. geophylla; red staining becoming very marked with age, particularly on the stipe which may be slightly swollen (sub-bulbous) at base; uncommon but not rare . . . . I. whitei
Cap to 50 mm, becoming coarsely fibrillose with age, sometimes entirely reddening thus similar to I. godeyi; gills beige then becoming spotted brick-red; stipe to 80 × 0.6(-1.0) mm; flesh whitish becoming reddish near surface; smell spermatic; spores smooth, amygdaliform with rounded apex, to 9.5 × 5.5 µm; hymenial cystidia metuloid, broadly lageniform, to 60(70) × 22 µm, walls to 2.5 µm thick; misshapen caulocystidia in top third of stipe as hymenial cystidia with cauloparacystidia amongst them; fruiting in autumn. Superficially easily confused with I. godeyi when mature, but the entirely pruinose and bulbous stipe together with earlier fruiting of the latter should suffice to separate the two species. (Figs 12, 14)
11
11*
12
Cap ± scaly, mid greyish brown or darkish brown; stipe often becoming darker at base; smell either strong: fruity and of peardrops, or complex: of stables / urine, sometimes with sweet fruity component on drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Cap ± smooth, tomentose to fibrillose, developing at most a few scales with age; either pale yellow-brown or whitish grey in which case developing greenish tinge at centre; smell always fruity, of peardrops, becoming intense and pervasive when contained. . . . . . . . . . . . . . . . . . . . . 13 Cap evenly coloured, mid greyish brown becoming dirty greenish or pinkish tinged, developing small appressed scales; cap flesh and stipe staining mildly pinkish red when exposed to air; smell complex, of stables/urine, when drying ± sweetish with fruity component . . . . . . . . . I. haemacta Fruitbody of medium (to large) stature; cap to 60(70) mm, rounded then flattened, sometimes with umbo, becoming coarsely fibrillose to scaly with age, pinkish where damaged; gills pale beige then darker with pink tinge where damaged; stipe to 60(80) × 8 mm, base ± bulbous, pale olivaceous and striate above, greenish grey with pinkish tinge and ± grooved below; spores smooth, amygdaliform with ± conical apex, to 11(12) × 6.5(7) µm (in one collection to 17 µm long with 2-sp. basidia); hymenial cystidia metuloid, cylindrical to lageniform, apex often notably narrow, to 65(80) × 20 µm, walls to 2 µm thick; caulocystidia either absent or present at stipe apex (more irregular than the hymenial cystidia); occasional on clayey calcareous soil under deciduous trees, often Corylus. (Figs 5, 7)
12*
Cap developing large dark brown scales on a paler cream ground, centre almost black; flesh infrequently weakly reddening; smell strong and fruity, of peardrops, pervasive when contained . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. erinaceomorpha Fruitbody of medium stature; cap to 50mm, margin persistently and angularly inrolled, then expanded with rounded umbo; gills unremarkable; stipe to 50 × 6mm, fibrillose, whitish above, beige below where sometimes mildly reddening, base occasionally dirty greenish; flesh whitish; spores smooth, amygdaliform with ± conical apex, to 10 × 6 µm; hymenial cystidia metuloid, often infrequent, cylindrical to subfusiform, to 65 × 20(25) µm, walls to 2 µm thick; caulocystidia absent; rare on calcareous soil under Fagus (and reportedly Quercus). (Figs 9, 10)
13
Cap whitish grey often with greenish tinge at centre, which long remains obtusely umbonate, finely fibrillose to tomentose; stipe sometimes dirty greenish grey at base; flesh infrequently weakly reddening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. corydalina Fruitbody of medium to large stature; cap to 60(80) mm, long remaining conical-campanulate with prominent rounded umbo; gills unremarkable; stipe to 80 × 10 mm, fibrillose, whitish above; spores smooth, amygdaliform with ± conical apex, to 9.5 × 6 µm; hymenial cystidia metuloid, often infrequent, ± cylindrical, rather thickset, to 65 × 20 (25) µm, walls to 1.5(2) µm thick; caulocystidia absent; occasional under deciduous trees, especially Fagus. (Figs 6, 8)
13*
Cap pale yellow becoming darker ochre-brown with age, when expanded usually losing umbo; and becoming ± flattened; stipe base not greenish grey; all parts ± reddening with age, damage or handling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
34
E L F GIH 14
Fruitbody of medium stature, cap to 50 mm, all parts (usually) reddening slightly to moderately. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. fraudans Cap pale yellow (at first whitish from veil) becoming darker ochre-brown with age, margin sometimes angularly inrolled when young, ± smooth then more roughened with age; gills unremarkable; stipe to 80 × 10 cm; smell pervasive, of peardrops; flesh tending to redden when exposed to air; spores smooth, amygdaliform with slight suprahilar depression, to 11(12) × 7.5 µm; hymenial cystidia metuloid, often infrequent, ± cylindrical, rather thickset, to 70 × 20 (25) µm, walls to 2.5(3)µm thick; caulocystidia absent; occasional under deciduous trees, especially Fagus. (Figs 1, 3)
14*
Fruitbody larger, to 70 mm or more, sometimes becoming misshapen, all parts reddening strongly; in other respects similar to I. fraudans; known also to occur with Helianthemum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. incarnata Morphological delimitation between I. incarnata and I. fraudans is not yet clear though DNA (Jacobsson & Larsson, 2012) has shown them to be distinct. It is possible that I. incarnata is the commoner of the two in Britain. One collection of I. incarnata growing with Helianthemum was observed to blacken after reddening as in some Russula species; it is not known if this is significant. It is also worth noting that there is a further alpine variety recognised in Scandinavia – I. fraudans var. chamaesalicis which occurs with Salix and Dryas. It is not improbable that our species with Helianthemum may prove to be different again once DNA work is carried out, particularly bearing in mind that this is now known to be the case with certain species of Amanita, Hebeloma, Cortinarius and Tricholoma associating with that host. (Figs 2, 4)
References Bandini, D. www.inocybe.org. An excellent website with descriptions and photos of most European species. Ferrari, E. (2006). Inocybe alpine e subalpine. Fungi non delineati 34–36. Edizioni Candusso. Ferrari, E. (2010). Inocybe dai litorali alla zona alpina. Fungi non delineati 54–55. Edizioni Candusso. Kuyper, T.W. (1986). A revision of the genus Inocybe in Europe. Persoonia Suppl. Vol. 3: 1–247. Leiden. Stangl, J. (2011). The genus Inocybe in Bavaria. (English Edit.) Publ. P. Cullington, A. McAdam et al.
Jacobsson, S. & Larsson, E. (2012). Inocybe in: Funga Nordica. Nordsvamp. Larsson, E. (2009). Taxonomy and evolutionary relationships within the species of section Rimosae (Inocybe) based on ITS,LSU and mtSSU sequence data. Persoonia 23: 86–98. Outen, A.R & Cullington, P. (2015). Keys to British species of Inocybe 3.1. www.bucksfungusgroup.org.uk
Further Reading Alessio, C.L. (1980). Iconographia Mycologica Vol. 29 suppl. 3. Inocybe.Trento, Italy.
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CBIB Update Six now available
Approximately 247 net additions (291 additions and 44 exclusions), almost all at species level, have been made to the Checklist since the publication of the book by Legon & Henrici in 2005 yielding an average of ca. 25 net additions per annum throughout the last decade.
A sixth update (March 2015) for the online Checklist of the British & Irish Basidiomycota (CBIB) is now available to download from the CBIB website: (http://www.basidiochecklist.info/LatestUpdates. asp) and P.M. Kirk has incorporated all the changes in the online database. As this is the first update since 2011, priority has been accorded to new additions and exclusions rather than to taxonomic and nomenclatural changes.
A. Martyn Ainsworth
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A key to reddening species of Inocybe in Britain >LL @
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hen working through a dichotomous key to a genus having a large number of species such as Inocybe, it can be somewhat frustrating to have to negotiate a long and often tortuous journey when one’s specimen has a particularly striking character. It can be tantalising to find no reference to this character along the way until perhaps just a brief mention at journey’s end. Mind you, having dabbled to some extent with devising such keys I do have sympathies for the challenged designer as well as the sometimes sorely tested user. It was with this issue in mind that I decided to design a key to the species of Inocybe that redden when bruised.
Subgenus Inosperma Section Rimosae (to which these two species belong) probably contain some degree of muscarin. I. erubescens, another reddening species of subgenus Inosperma, has sufficient quantities of muscarin to render it deadly poisonous. Bearing in mind the considerable danger of consuming I. erubescens in error, it makes good sense for those of us who enjoy collecting for the pot never to eat (or even taste) any member of this genus. In any case, taste is not amongst the characters used in identification in Inocybe. Microscopy It is always essential to observe microscopic characters when working with this genus, and I make no apology for referring to spore shape and cystidia characteristics right from the start of the key. If this is not your forte then read no further – making reliable determinations of Inocybe without recourse to a microscope is unfortunately not possible. For examination the only chemical needed is 10% ammonia, no other stains are necessary, and hymenial cystidia (those found on the gill) are quite easy to locate compared to some other genera. Another plus: the fruitbodies as a rule are quite resilient and will normally survive for several days in the fridge or even in the post, the only proviso being that as with most agarics a sporeprint for really accurate spore measurement needs to be acquired from an unrefrigerated specimen. For the purposes of this key, only cheilocystidia (found on the gill edge) need be examined: if these are thin-walled one can safely assume that pleurocystidia (found on the gill face) are absent; conversely if thick-walled and capped with crystals (metuloids) one can safely assume that pleurocystidia are also present (if both sorts are absent on the gill edge then one probably has a different genus).
Types of reddening Of the approximately 140 species of Inocybe in Britain, 15 show reddening in some part of the fruitbody to varying degrees. The reddening itself varies from species to species and can be bright red, vinaceous, orange-red to rust-red: it can occur in some species throughout the fruitbody, in others just in the stipe. In two species, I. corydalina and I. erinaceomorpha, reddening is only rarely present in my experience, but they are included here for completeness. In all but two, I. adaequata and I. rhodiola, where the dark vinaceous red staining found in the cap, stipe and flesh seems to be inherent from the start, the reddening is not present in young, pristine specimens but develops either as they mature or with handling or damage. Toxins Though many species of Inocybe (including some of the reddening species) are reported to contain dangerously poisonous toxins, including psilocybin and muscarin, Kuyper (1986) comments that one of the reddening species, I. adaequata [therefore possibly I. rhodiola also?], contains no muscarin and is considered the only safely edible member of the genus. All other members of
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E L F GIH Caulocystidia Stipe pruinosity and locating caulocystidia correctly is generally much harder to assess and (in my experience) is the commonest cause of errors and confusion in identification of Inocybe, though only the second half of this key requires this information. I have found that even with the best dissecting scope it is not always possible to judge safely whether pruinosity apparent on the stipe is due to true caulocystidia or merely hairs (surface hyphae which become detached to some degree). The only safe way is to take the finest strip of fibre from the stipe and examine it under a compound microscope in ammonia, at the same time noting from exactly which part of the stipe your strip came. To use the key you may need to know whether caulocystidia are: 1) absent even at the apex, 2) present only at the apex, or 3) present in the top 3rd or in one species right down to the base. A word of caution: it is all too easy to remove or damage caulocystidia through overhandling on collection, thus giving you a false reading when assessing with a microscope. An added benefit to making up a slide from the stipe is that the spores found there will have dropped naturally and will therefore be mature. Thus in the absence of a spore print this gives you more accurate measurements than if measured from a gill squash.
included therein, also that I. rhodiola though included is not there accepted as British. This is rectified in Edition 3 (Outen & Cullington, 2015), now available online. Both species are recognised by some (if not all) authorities in Europe and have now been found in Britain. At present with this genus we are reliant upon workers abroad whose research using DNA is proving extremely useful. Until such time as British collections of Inocybe can be similarly researched there will inevitably remain an element of doubt over declaring new species here. Species descriptions A fairly detailed description follows each species as it occurs in the key. Please note, however, that to avoid unnecessary repetition the main characters already covered in the relevant couplet are not then duplicated in the description unless further explanation is useful. Stature (medium, large etc) is relative to members of the genus as a whole. Inocybe Key For complete British coverage of this genus there is now a new edition of Outen and Cullington’s Keys to British species of Inocybe (Outen & Cullington, 2015), available free of charge online, hosted by the Bucks Fungus Group’s website: (www.bucksfungusgroup.org.uk). Edition 3:1 April 2015 (Cullington) is much changed from previous editions with, amongst other things, a complete revision of Key D for nodulose-spored marginate species, also the key to reddening species included in this article. This latest version is recommended in place of previous editions. To gain access you will need to email me for the necessary code: ([email protected]).
Measurements of cystidia and spores With this genus the size and shape of cystidia (and to a lesser extent spores as well) can vary far more than is met with in some other genera even within one gill, let alone one specimen. Re size: I find the best approach is to make note of the longest / widest / thickest-walled examples rather than attempt to assess an average, then compare this with the greatest measurement given in a description (bearing in mind that cystidia found on the gill face tend to be slightly bigger than those on the edge). The same advice applies to measuring spores. Regarding cystidia shape: it is well-nigh impossible to cover in a nutshell the variety of shapes which may occur in one gill preparation; I advise aiming for an overall general impression, taking more note of the larger examples which tend to be more typical.
All photographs of fruitbodies and microscopy are © Penny Cullington unless otherwise attributed.
Two new species Owners of the printed versions of Outen & Cullington Keys will note that I. armeniaca is not
/3
E L F GIH .+0 -0. . , 0 .(,. 1
Spores nodulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. bresadolae Fruitbody of medium stature; cap to 45(50) mm, campanulate then flat with an umbo, tomentose to ± scalyfibrillose, pale ochraceous brown with orange tint, reddening near margin; gills beige, eventually reddening; stipe to 45 × 7 mm, yellowish, sometimes reddening, sometimes with (marginate) bulb, entirely pruinose with caulocystidia; flesh with guaiac stains intense wine-red; smell sweetish as in I. bongardii; spores 7–10 × 5–7 µm with a few small nodules; hymenial cystidia metuloid, fusiform, to 75 × 17 µm, walls to 3 µm thick, yellow in ammonia; under deciduous trees in summer, reportedly rarely in autumn. Rare: very few British records. (Figs 27, 28).
1*
Spores smooth – either ellipsoid to phaseoliform or amygdaliform . . . . . . . . . . . . . . . . . . . . . . . . 2
2 2*
Spores ellipsoid to phaseoliform; cheilocystidia thin-walled, pleurocystidia absent . . . . . . . . . . . 3 Spores amygdaliform; cheilocystidia thick-walled, mostly crystal-tipped, pleurocystidia present 8
3
Cap predominantly scaly, squamulose to squarrulose, some shade of brown; rust-red staining confined to stipe where handled or damaged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Cap smooth to markedly fibrillose, never becoming predominantly scaly, either whitish or vinaceous brown; reddening occurring throughout fruitbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3*
4
Cap scales darkish brown on a paler brown ground, becoming ± squarrulose at centre; smell strong and disagreeable, earthy or musty; mainly under Picea . . . . . . . . . . . . . . . . . I. cervicolor
Fruitbody of medium stature; cap to 45 mm, ± domed then flat; gills pale yellowish then brown; stipe to 100 x 6 mm, concolorous, ± flocculose above, streaky fibrillose below where becoming rust-red when handled; spores smooth, ellipsoid to subphaseoliform, 11–15 × 6.5–8.5 µm; cheilocystidia thin-walled, wavycylindrical to slenderly clavate, to 50(60) × 14(16) µm, pleurocystidia absent. Uncommon. (Figs 25, 26).
4*
Cap scales mid-brown on a paler buff to pinkish brown ground, remaining ± squamulose; smell either pleasant and fruity, or if unpleasant then fishy; mainly under Fagus and Quercus . . . . . . 5
5
Smell sweetish with flowery or fruity component. . . . . . . . . . . . . . . I. bongardii var. bongardii
Fruitbody of medium stature though stipe sometimes disproportionately long; cap to 50(60) mm, ± campanulate; gills unremarkable greyish brown; stipe to 80(90) × 9 mm, concolorous, ± flocculose above, streaky fibrillose below where becoming rust-red when handled; smell usually pleasant (though when found with Helianthemum smell faint, ± earthy?); spores smooth, ellipsoid to subphaseoliform, 11–16 × 6.5–9 µm; cheilocystidia thin-walled, wavy-cylindrical to slenderly clavate, to 60 × 18 µm, pleurocystidia absent; under Fagus, Quercus and Castanea, also with Helianthemum; occasional. (Figs 22, 24).
5*
Smell unpleasant, slightly fishy cf. Macrocystidia cucumis . . . . . . I. bongardii var. pisciodora Details as for the type variety above, but smell complex: reportedly sometimes with a Pelargonium component and at some stage becoming unpleasant and fishy. It is advisable to contain specimens and check periodically after collection to assess the smell which can be elusive, even ephemeral. Rarely reported; I have found this variety under Fagus.
6
Cap smooth; cap and stipe initially white, soon becoming ochraceous to flesh-coloured, developing bright red stains with age and/or damage; smell ± sickly, of soap; fruiting in June or July under Fagus on calcareous soil. Dangerously poisonous. . . . . . . . . . . . . . . . . . . . . . . . . . . . I. erubescens Fruitbody of medium to large stature; cap to 70 mm or more, at first with abruptly downturned margin, then with obtuse umbo, splitting with age; gills pale grey-beige then dirty olivaceous, bruising red where touched; stipe to 70(90) × 15(20) mm, concolorous, sometimes with clavate base; flesh off-white to pinkish, bruising red; spores smooth, ellipsoid to subphaseoliform, to 14 × 7.5 µm; cheilocystidia slightly clavate, to 65 × 15(20) µm, pleurocystidia absent; uncommon (Figs 21, 23).
6*
Cap markedly streaky fibrillose, cap and lower stipe dirty russet to vinaceous brown, upper stipe remaining paler; flesh discolouring pinkish to vinaceous from the start; smell different; fruiting in autumn under various deciduous trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
/1
E L F GIH 7
Fruitbody robust, cap to 60(80)mm with obtuse umbo and stipe to 70 x 12 µm; maturing dirty brownish purple; smell unpleasant, earthy or of beetroot; not uncommon especially under Quercus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. adaequata
Cap streaky fibrillose, sometimes furrowed, dirty vinaceous red-brown; gills at first pale beige then darker with reddish tinge; stipe ± cylindrical, not pruinose, not bulbous, pale at first ( almost white at apex), then from the base increasingly pinkish to dark reddish brown; stipe and cap flesh white then soon pinkish, becoming darker vinaceous brown from stipe base upwards; spores smooth, ellipsoid to slightly phaseoliform, 11–13(14) × (5)6–7(8) µm; cheilocystidia thin-walled, wavy-cylindrical, to 70 × 15 µm; under deciduous trees, usually Quercus; occasional. Delimitation between I. adaequata and the smaller but similar I. rhodiola (shown to be distinct with DNA, Larsson 2009) is not yet clear, although I. rhodiola appears to retain its pink stipe tones rather than darkening to vinaceous brown; several British collections matching well to I. rhodiola have been made since then. (Figs 17, 18).
7*
Fruitbody smaller, more delicate and slender, umbo acute to obtuse; stipe usually pinkish below; smell faint; probably rare, found so far under Fagus in S. England . . . . . . . . . . . . . . . I. rhodiola Microscopic details are as far as known similar to I. adaequata. See comments re delimitation under that species. (Figs 19, 20).
8
Stipe entirely pruinose with caulocystidia, base marginately bulbous; cap pale yellowish but fruitbody becoming increasingly stained bright red, sometimes ± entirely so when old . . I. godeyi Fruitbody of small to medium stature; cap to 40(50) mm, conical then with ± acute umbo, finely fibrillose, sometimes scaly with age, almost white then ochraceous but soon strongly (even entirely) reddening; gills pale beige then ochraceous, reddening; stipe to 40(50) × 6 mm, concolorous, reddening; flesh pale but reddening; smell acidic; spores smooth, amygdaliform, to 12 × 7 µm; hymenial cystidia metuloid, swollen fusiform, to 65(70) × 25 µm, walls to 1.5(2) µm thick; under deciduous trees on calcareous soil; occasional, tending to fruit in summer to early autumn. (Figs 11, 13).
8*
Stipe either pruinose with caulocystidia descending no lower than the top third, or caulocystidia virtually absent though may appear pruinose/hairy at apex, base not bulbous . . . . . . . . . . . . . . . 9
9.
Cap white and ± silky smooth, developing orange-red staining as it matures, especially where handled; stipe white with tendency to redden, pruinose with caulocystidia in top 3rd (sometimes less far down?); smell unremarkable, spermatic; spores with ± obtuse apex; under conifers and deciduous trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Cap some other colour and not silky smooth, either fibrillose or scaly; stipe appearing pruinose at apex but caulocystidia virtually absent; smell unusual: often strong, fruity, of pear-drops or of urine/stables; spores with ± conical apex; only under deciduous trees . . . . . . . . . . . . . . . . . . . . . 11
9*
10
Fruitbody small and slender, similar in appearance to I. geophylla but additionally with orangered staining developing on part of cap and / or stipe. Rare. . . . . . . . . . . . . . . . . . . . . I. armeniaca
Cap to 20(30) mm, silky smooth to finely fibrillose, conical usually with acute umbo when young, white to palest cream but developing orange-red patches which separate it from I. geophylla in the field; gills unremarkable, possibly also orange-red where damaged; stipe to 50 × 2(5) mm, ± cylindrical, pruinose near apex, cortina present when immature, not bulbous, concolorous, possibly becoming orange-red with age; flesh whitish, also possibly becoming orange red with age; smell if present spermatic; spores smooth, amygdaliform with rounded apex, av. 9 × 5 µm; hymenial cystidia metuloid, similar to those of I. geophylla, walls to 2 µm thick, pale yellow in ammonia; misshapen caulocystidia present near stipe apex amongst cauloparacystidia which extend further down (though according to Bon caulocystidia absent?). Specific tree association in Britain is as yet not established; in Europe reported from both sandy and calcareous soil under a mix of conifers and deciduous trees. I have seen two collections in Britain (see Field Mycology Vol 15(4): 134) though sadly the material was not retained. Further collections are needed to clarify species details and provide fungarium specimens to back up its presence here. (Figs 15, 16).
32
E L F GIH 10*
Fruitbody larger, of medium stature, cap to 40 mm or more, ± domed, rarely with acute umbo, i.e. not reminiscent of I. geophylla; red staining becoming very marked with age, particularly on the stipe which may be slightly swollen (sub-bulbous) at base; uncommon but not rare . . . . I. whitei
Cap to 50 mm, becoming coarsely fibrillose with age, sometimes entirely reddening thus similar to I. godeyi; gills beige then becoming spotted brick-red; stipe to 80 × 0.6(-1.0) mm; flesh whitish becoming reddish near surface; smell spermatic; spores smooth, amygdaliform with rounded apex, to 9.5 × 5.5 µm; hymenial cystidia metuloid, broadly lageniform, to 60(70) × 22 µm, walls to 2.5 µm thick; misshapen caulocystidia in top third of stipe as hymenial cystidia with cauloparacystidia amongst them; fruiting in autumn. Superficially easily confused with I. godeyi when mature, but the entirely pruinose and bulbous stipe together with earlier fruiting of the latter should suffice to separate the two species. (Figs 12, 14)
11
11*
12
Cap ± scaly, mid greyish brown or darkish brown; stipe often becoming darker at base; smell either strong: fruity and of peardrops, or complex: of stables / urine, sometimes with sweet fruity component on drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Cap ± smooth, tomentose to fibrillose, developing at most a few scales with age; either pale yellow-brown or whitish grey in which case developing greenish tinge at centre; smell always fruity, of peardrops, becoming intense and pervasive when contained. . . . . . . . . . . . . . . . . . . . . 13 Cap evenly coloured, mid greyish brown becoming dirty greenish or pinkish tinged, developing small appressed scales; cap flesh and stipe staining mildly pinkish red when exposed to air; smell complex, of stables/urine, when drying ± sweetish with fruity component . . . . . . . . . I. haemacta Fruitbody of medium (to large) stature; cap to 60(70) mm, rounded then flattened, sometimes with umbo, becoming coarsely fibrillose to scaly with age, pinkish where damaged; gills pale beige then darker with pink tinge where damaged; stipe to 60(80) × 8 mm, base ± bulbous, pale olivaceous and striate above, greenish grey with pinkish tinge and ± grooved below; spores smooth, amygdaliform with ± conical apex, to 11(12) × 6.5(7) µm (in one collection to 17 µm long with 2-sp. basidia); hymenial cystidia metuloid, cylindrical to lageniform, apex often notably narrow, to 65(80) × 20 µm, walls to 2 µm thick; caulocystidia either absent or present at stipe apex (more irregular than the hymenial cystidia); occasional on clayey calcareous soil under deciduous trees, often Corylus. (Figs 5, 7)
12*
Cap developing large dark brown scales on a paler cream ground, centre almost black; flesh infrequently weakly reddening; smell strong and fruity, of peardrops, pervasive when contained . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. erinaceomorpha Fruitbody of medium stature; cap to 50mm, margin persistently and angularly inrolled, then expanded with rounded umbo; gills unremarkable; stipe to 50 × 6mm, fibrillose, whitish above, beige below where sometimes mildly reddening, base occasionally dirty greenish; flesh whitish; spores smooth, amygdaliform with ± conical apex, to 10 × 6 µm; hymenial cystidia metuloid, often infrequent, cylindrical to subfusiform, to 65 × 20(25) µm, walls to 2 µm thick; caulocystidia absent; rare on calcareous soil under Fagus (and reportedly Quercus). (Figs 9, 10)
13
Cap whitish grey often with greenish tinge at centre, which long remains obtusely umbonate, finely fibrillose to tomentose; stipe sometimes dirty greenish grey at base; flesh infrequently weakly reddening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. corydalina Fruitbody of medium to large stature; cap to 60(80) mm, long remaining conical-campanulate with prominent rounded umbo; gills unremarkable; stipe to 80 × 10 mm, fibrillose, whitish above; spores smooth, amygdaliform with ± conical apex, to 9.5 × 6 µm; hymenial cystidia metuloid, often infrequent, ± cylindrical, rather thickset, to 65 × 20 (25) µm, walls to 1.5(2) µm thick; caulocystidia absent; occasional under deciduous trees, especially Fagus. (Figs 6, 8)
13*
Cap pale yellow becoming darker ochre-brown with age, when expanded usually losing umbo; and becoming ± flattened; stipe base not greenish grey; all parts ± reddening with age, damage or handling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
34
E L F GIH 14
Fruitbody of medium stature, cap to 50 mm, all parts (usually) reddening slightly to moderately. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. fraudans Cap pale yellow (at first whitish from veil) becoming darker ochre-brown with age, margin sometimes angularly inrolled when young, ± smooth then more roughened with age; gills unremarkable; stipe to 80 × 10 cm; smell pervasive, of peardrops; flesh tending to redden when exposed to air; spores smooth, amygdaliform with slight suprahilar depression, to 11(12) × 7.5 µm; hymenial cystidia metuloid, often infrequent, ± cylindrical, rather thickset, to 70 × 20 (25) µm, walls to 2.5(3)µm thick; caulocystidia absent; occasional under deciduous trees, especially Fagus. (Figs 1, 3)
14*
Fruitbody larger, to 70 mm or more, sometimes becoming misshapen, all parts reddening strongly; in other respects similar to I. fraudans; known also to occur with Helianthemum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. incarnata Morphological delimitation between I. incarnata and I. fraudans is not yet clear though DNA (Jacobsson & Larsson, 2012) has shown them to be distinct. It is possible that I. incarnata is the commoner of the two in Britain. One collection of I. incarnata growing with Helianthemum was observed to blacken after reddening as in some Russula species; it is not known if this is significant. It is also worth noting that there is a further alpine variety recognised in Scandinavia – I. fraudans var. chamaesalicis which occurs with Salix and Dryas. It is not improbable that our species with Helianthemum may prove to be different again once DNA work is carried out, particularly bearing in mind that this is now known to be the case with certain species of Amanita, Hebeloma, Cortinarius and Tricholoma associating with that host. (Figs 2, 4)
References Bandini, D. www.inocybe.org. An excellent website with descriptions and photos of most European species. Ferrari, E. (2006). Inocybe alpine e subalpine. Fungi non delineati 34–36. Edizioni Candusso. Ferrari, E. (2010). Inocybe dai litorali alla zona alpina. Fungi non delineati 54–55. Edizioni Candusso. Kuyper, T.W. (1986). A revision of the genus Inocybe in Europe. Persoonia Suppl. Vol. 3: 1–247. Leiden. Stangl, J. (2011). The genus Inocybe in Bavaria. (English Edit.) Publ. P. Cullington, A. McAdam et al.
Jacobsson, S. & Larsson, E. (2012). Inocybe in: Funga Nordica. Nordsvamp. Larsson, E. (2009). Taxonomy and evolutionary relationships within the species of section Rimosae (Inocybe) based on ITS,LSU and mtSSU sequence data. Persoonia 23: 86–98. Outen, A.R & Cullington, P. (2015). Keys to British species of Inocybe 3.1. www.bucksfungusgroup.org.uk
Further Reading Alessio, C.L. (1980). Iconographia Mycologica Vol. 29 suppl. 3. Inocybe.Trento, Italy.
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CBIB Update Six now available
Approximately 247 net additions (291 additions and 44 exclusions), almost all at species level, have been made to the Checklist since the publication of the book by Legon & Henrici in 2005 yielding an average of ca. 25 net additions per annum throughout the last decade.
A sixth update (March 2015) for the online Checklist of the British & Irish Basidiomycota (CBIB) is now available to download from the CBIB website: (http://www.basidiochecklist.info/LatestUpdates. asp) and P.M. Kirk has incorporated all the changes in the online database. As this is the first update since 2011, priority has been accorded to new additions and exclusions rather than to taxonomic and nomenclatural changes.
A. Martyn Ainsworth
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