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A field key to British non-resupinate hydnoid fungi
Field Mycology Volume 1(3), July 2000
A FIELD KEY TO BRITISH NON-RESUPINATE HYDNOID FUNGI Gordon Dickson Flagstones, 72 Catisfield Lane, Fareham, Hants PO15 5NS
T
he key employs characters discernible in the field. With two exceptions, all species have been examined in the fresh condition: the exceptions are Bankera violascens (of which there has been only one British record) and Hydnellum auratile. Hydnellum ferrugineum has not been seen in active growth. In the past, insufficient attention has been paid to the tree associates. The associated trees given are the ones under which I, personally, have found the species.Where there could be doubt due to the presence of more than one tree species within root range I have not drawn any conclusion.The term ‘mixed woodland’ has no meaning in this context as the fungus is mycorrhizal with only one tree species at a time. It must also be remembered that in different countries different host trees are found. This key refers only to Britain. In order to reinforce the field identification, some microscopic features have been appended. KEY TO GENERA
1) -
Growing on pine cones, stipe thin, tall, excentric . . . . . . . . . . . . . . . . . . . . .Auriscalpium vulgare Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2(1) Growing on wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 Growing on the ground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 3(2) Jelly-like, under 5 cm; dark grey to brown or fawn upper surface, pale grey spines. On conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudohydnum gelatinosum Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Hydnellum scrobiculatum with beautifully zonate and radially wrinkled caps, which may fuse together in large masses. Photograph © Gordon Dickson 99
Field Mycology Volume 1(3), July 2000 4(3)Over 5 cm diameter, white or cream, rubbery consistency; on broad-leaf trees, usually beech . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium see Key A Bracket with zoned pileus very similar in appearance to Stereum hirsutum but with buff or orange spines on underside; on hardwoods, usually beech . . . . . . . . . . . . . . . . . .Steccherinum ochraceum 5(2) White, friable, very small spines, on soil, moss or pine needles, smelling of iodoform when fresh; shape very variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sistotrema confluens Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 6(5) Cap fleshy ie. Agaricus consistency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7 Fibrous or corky consistency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 7(6) Spines more or less colour of cap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum see Key E Spines a different colour from cap surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8 8(7) Spines grey; spores brown; cap coloured. . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon see Key F Spines white to buff; spores white; cap white, then brown . . . . . . . . . . . . . . . .Bankera see Key C 9(6) Spines (and spores) white; cap black, blue-black to brown . . . . . . . . . . . . . . .Phellodon see Key D Spines (and spores) brown; cap some shade of brown +/- blue tints . . . . . . .Hydnellum see Key B Key A - HERICIUM 1 Mature spines > 3 cm. long, hanging; usually high on beech (until the tree falls!), recorded once on oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium erinaceum Mature spines < 3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 2(1) Lobed; upper surface strigose (bristly-hairy); on beech . . . . . .Hericium(= Creolophus) cirrhatum Whole fungus consisting of a mass of intertwining branches with short spines falling comb-like along their length; on beech . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium coralloides Key B - HYDNELLUM 1) Cap whitish, pink, buff or fulvous, darkening, consisting of fused pileoli, velvety at first; darkening with age, “lumpy” but not wrinkled or markedly zoned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Cap brown with/without blue tints; wrinkled and zoned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2(1) Cap at first fulvous or buff, velvety. Later the tomentum collapses and the surface darkens; with oak or chestnut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum spongiosipes Cap surface at first whitish to pink, “lumpy” and showing fused pileoli, then darkening; in active growth may show red guttation drops; taste mild, or mealy; with pine . . . .Hydnellum ferrugineum 3(1) Cap margin, spines and flesh all showing blue colours . . . . . . . . . . . . . . . . .Hydnellum caeruleum Cap brown (no blue), surface radially wrinkled and concentrically zoned . . . . . . . . . . . . . . . . . .4 4(3) Cap flesh thin (< 2mm), almost papery; with pine . . . . . . . . . . . . . . . . .Hydnellum scrobiculatum Not this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 5(4) Cap shades of brown, thin-fleshed, often tuberculate in the centre; with oak or chestnut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum concrescens Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 6(5) Cap flesh thick (>3mm.); surface brown, rough and with growing edge often pink with red droplets; taste intensely hot, often after a delay; with pine . . . . . . . . . . . . . . . . . .Hydnellum peckii
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Taste not hot, cap with some orange tints; flesh thick; with pine . . . . . . . . . . . . . . . . . . . . . . . . .7
7(6) Stem orange (or partly so), flesh brown tinted orange in stem only . . . . . .Hydnellum aurantiacum Stem brown, pileus flesh brown, stem flesh dull orange with extensions through to pileus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum auratile NB. See species notes before recording species in couplets 2, 4 or 7. KEY C - BANKERA 1 Cap and stem whitish with slight violet tint; spines pale grey; smell sweetish when fresh, of curry when dry, with spruce (only one record to date) . . . . . . . . . . . . . . . . . . . . . . . .Bankera violascens Cap surface buff or cinnamon buff; spines and stem white, then cream; with pine, often under Calluna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bankera fuligineoalba KEY D - PHELLODON 1 Stipe black, pileus some shade of “black” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Stipe and pileus some shade of brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2(1) Cap zoned with a white edge, centre commencing brown and becoming brown-black, pileoli usually fused, not overlapping, with pine, spruce or birch . . . . . . . . . . . . . . .Phellodon melaleucus Cap with sharply defined white edge, centre with blue-black silky tomentum without shades of brown. Pileoli usually overlapping, with oak in the south, pine or spruce in Scotland. (Photo on back cover) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phellodon niger 3(1) Multiple, zoned pileoli, usually fused, with pine or spruce . . . . . . . . . . . . . . .Phellodon tomentosus Cap buff colour greying with age, not or only slightly zoned, stipe tomentose, with broad-leaf trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phellodon confluens KEY E - HYDNUM 1 Cap usually fairly robust (over 5 cm) pale cinnamon to almost cream; spines almost concolorous, decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum repandum Cap cinnamon to orange, not usually over 5cm; spines concolorous, free or adnate, NOT decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum rufescens KEY F - SARCODON 1 Cap with recurved scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Cap smooth, cracking in central area but without recurved scales; stem base blue-grey, both externally and in flesh; with pine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon glaucopus 2(1) Cap scaly, stem pale except at base which is blue-grey internally and externally; with sweet chestnut or oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon scabrosus Cap with dark erect scales on a pale ground; stem pale throughout its length. With pine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon squamosus
Notes on the species in the key Some of the Hydnellum and Phellodon species are relatively easy to recognise when they are in active growth but extremely difficult when growth has ceased, even using microscopic methods. Fortunately many have an extended season and it is often possible to find a younger specimen than the old one first located. It is not worth spending time on old material. I have noted here the trees under which I have found the species. Where more than one tree has been noted within root range I have recorded this (the host tree is not always the nearest one). If the same species of tree occurs repeatedly and is never absent there is a strong probability that this is the host species. For the present I have indicated the tree which I 101
Field Mycology Volume 1(3), July 2000 am confident is the mycorrhizal partner in Britain and have shown in square brackets the species indicated by Pegler et al.(1997) for Europe. Baird(1985) quotes trees within 20m from the fungus but this is an uncertain method and I believe that it is better to use for reference those instances where there is irrefutable proof that no other species of tree is involved, even though this limits the number of observations. I would be grateful to have further information on new determinations in order to upgrade this key. In particular I would like to have notes of fungi found in pure stands of one species of tree (being careful to note the odd birch which sneaks in!) For the present it seems sufficient to note the genus of tree but if the species can be named so much the better. Auriscalpium vulgare The ‘ear-pick’ fungus cannot be confused with anything else. Always on the cones of pine (sometimes buried). Bankera fuligineoalba The combination of white stem and spines, with a variably brownish pileus surface and a ‘mushroom’ consistency leaves no doubt as to its identity. It has a very weak aroma when fresh but smells quite strongly of curry when dry; it occurs with pine and seems to prefer fruiting under the overhang of heather, especially along paths [pine]. Bankera violascens This species has been found only once (September 1996) by a track through a spruce plantation near Loch Morlich in Scotland. The spines are white but the rest of the fungus is a dull violaceous colour. It has a sweet, not unpleasant smell when fresh. The one collection was at a trackside where it had already been damaged by passing boots. It occurs only with spruce.
Hericium species On close examination the intertwined growth of H. coralloides is distinctive and the two remaining species can be separated on spine length. Even from a distance H. erinaceum can be seen either to be a round ball or with horizontal tiers whereas H. cirrhatum consists of lobes protruding in any direction. Hydnellum aurantiacum and H.auratile I have not seen fresh material of Hydnellum auratile but have seen many collections of H.aurantiacum. The latter has a stem which is orange at least in parts, there is orange on the cap surface and certainly the brown flesh is suffused with orange in the stipe. H. auratile is stated to have a brown stem, very little orange on the cap surface and the orange flesh in the stipe extends up into the pileus. Both occur only with pine. Hydnellum caeruleum Unmistakable among British species of Hydnellum; the blue coloration during growth
Hydnellum caeruleum showing the blue, tomentose margin of actively growing material. Photograph © Gordon Dickson. 102
Field Mycology Volume 1(3), July 2000 marks it out immediately. When old this colour disappears from the surface and spines but is retained in the flesh. With Caledonian pine [pine, spruce, beech]. Hydnellum concrescens and H. scrobiculatum These species are lumped together because macroscopically their appearance overlaps so much that they are almost indistinguishable. Both have radially wrinkled and concentrically zoned brown pilei which may or may not be fused. Microscopically they can be distinguished by the tubercles on the spores which in H.concrescens have a double crest (like a two-humped camel) whilst in H.scrobiculatum they are single-humped. Also the spores are slightly larger (5-6.5 x 4-5μm as against 4.5-5.5 x 3-3.5μm for H. concrescens). In the field at present I have found H. concrescens to associate with oak or chestnut and H.scrobiculatum with pine. Hydnellum ferrugineum I have not seen material of this species when in active growth, only when fully mature. It is closely related to H.spongiosipes but is a more reddish brown colour and occurs with pine. It may show red ‘guttation drops’ when actively growing but can be distinguished immediately from H. peckii which also has red drops - by the mild taste and zoned flesh [pine, spruce]. Hydnellum peckii Dark brown and thick-fleshed, this species may have a lumpy or wrinkled pileus surface but it has one distinguishing character unique in British Hydnellum species. It has an intensely hot or acrid taste, felt, after a variable delay, at the back of the throat. With experience one learns to spit out before this stage is reached! I have found it only with Caledonian pine [pine, spruce]. Hydnellum spongiosipes This species is easy to recognise: its pileus surface can best be described as ‘lumpy’ and thickfleshed. It is not wrinkled and the colour changes with the collapse of the surface tomentum from clay-buff to brown vinaceous or sepia. The concrescent pileus shows little evidence of its separate origins but the multiple stems are clad in a soft, humus - binding base. I have found it only with oak, usually in moss banks [oak, beech, chestnut]. Hydnum repandum & H.rufescens There is little difficulty in identifying Hydnum in the field but separating the two species is less easy
especially as more or less intermediate forms exist. Apart from the colour difference, note should be taken of the distribution of the spines: those in H.repandum are decurrent, often reaching half way down the stem whilst those of H.rufescens are usually free of the stem or only just reaching it. It is also a less robust fungus. Oak, beech, birch. [oak, beech, birch, maple, hazel, pine, spruce]. Phellodon confluens The distinguishing feature of this species is its unzoned pileus surface. During growth it has the usual white pileus margin of all Phellodon species but otherwise the surface is a ferruginous buff, somewhat rough and slightly wrinkled. It is relatively thick fleshed, becoming top-shaped when fully grown and with a relatively short stipe, the spines extending to the level of the moss in which it usually grows. I have found it only with oak on clay [chestnut, oak, beech]. Phellodon melaleucus & P. niger Both these species are characterised by having a sharply defined white border to the fruit-body which consists of fused pilei about 2mm. thick tapering to almost paper-thin at the edge, often giving the appearance of a single, very large cap. The underside, however, reveals multiple stems. The white border is present only during active growth. During this phase they can be distinguished in the field by the character of the cap which catches the light as it is tilted in the hand and is a blue-black colour. P. melaleucus has a duller tomentum and the colour is brown-black. In the laboratory an additional check is the partly encrusted, greening, hyphae in the context of P. niger when mounted in alkali; those of P. melaleucus are olive-green to black. I have found P. melaleucus with pine, spruce and birch on sandy soil [broadleaf and coniferous trees]. P. niger I have found under oak on heavy marls in the New Forest and under Caledonian pine and planted spruce on sandy soils in Scotland [chestnut, beech & pine]. Phellodon tomentosus Unlike P. niger in which the pileoli lose their individuality on fusing, those of P. tomentosus retain a markedly zoned appearance in shades of brown, darker at the centre and, during growth, with a white margin. They are thin and papery at the edges especially when growth has ceased. This appearance is unmistakable in the field. Under pine [pine].
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Field Mycology Volume 1(3), July 2000 Pseudohydnum gelatinosum A lumpy, tongue-shaped mass of firm, translucent jelly, grey, brown or fawn on the surface, paler underneath. The only jelly-like spine-fungus. Always on dead conifer. Sarcodon glaucopus At a casual glance this species could be mistaken for an old specimen of S.imbricatus especially as it occurs in the same habitat, but it has no clamp connections, the pileus surface is darker and not scaly (sepia/umber), the spines grey and when it is inverted the lower part of the stem is seen to be grey. This colour is also present in the adjacent flesh. It is probably not very uncommon in some areas of Scotland, having been overlooked in the past. Caledonian pine [pine]. Sarcodon scabrosus Until recently this was known as S. imbricatus in Britain, but it is now known that these are distinct species and that the latter is not British (see Field Mycology 2: 42-43). This species is the only Sarcodon occurring with broad-leaf trees in Britain. It is usually with sweet chestnut or oak in Britain (though Fries’ original description says ‘In pinetis montanis raro’). The cap surface is chestnut brown, darkening with age, the scales being adpressed except in the centre; the spines are grey and the stem base also grey but it cannot be confused with the previous species as it has adpressed or slightly recurved scales and occurs with broad-leaf trees [oak, chestnut, pine]. Sarcodon squamosus The commonest of the Sarcodon species, this occurs from Scotland to the New Forest and is invariably associated with Scots pine in Britain. It has a ‘mushroom’ consistency and the scaly cap may vary from buff to a darker cigar brown in the centre. The scales may be adpressed in the outer part of the cap to upstanding in the centre. The flesh is whitish, the spines grey and the stem white. Like other spine fungi it seems to like to fruit under the protection of heather either at tracksides or beside rivers. Microscopically it can be differentiated from S. scabrosus by having clamp connections. Other Sarcodon species One other Sarcodon has been found in Britain: Sarcodon regalis was collected (and named) from Windsor Great Park in 1968: it is now regarded as extinct in Britain. Sarcodon fuligineoviolaceus, was listed as endangered by Ing (1992), but Pegler et al. (1997) were unable to find any evidence of it
having occurred in Britain. Sarcodon leucopus has not yet been recorded in Britain. It has a smooth, pale cap and could be mistaken for a Bankera but for the coloured spines; it occurs with conifer so should be looked for in Scotland. Sistotrema confluens This is not really a spine fungus but a poroid fungus with irpicoid pores. When growing on compacted soil it sometimes makes a stem and pileus and the pores can then be seen. Growing on a bank the hymenial surface can appear hydnoid. Its fragility and smell of iodoform when fresh identify it. Known chiefly but not exclusively from pines (it has been found with oak in the New Forest) but its ecology seems uncertain. Steccherinum ochraceum A fungus frequently missed because of its resemblance to Stereum hirsutum which is so common that the underside is left uninspected. The short spines on the underside are white, cream or orange. On deciduous wood. Acknowledgements I would like to thank P.D. Orton for many helpful comments on the first draft and for supplying me with (and translating) the Britzelmayr paper referred to below and to Martin Ainsworth and Adrian Newton for further helpful comments. Literature consulted Baird, R.E.(1986). Study of the stipitate hydnums from the Southern Appalachian Mountains Genera: Bankera, Hydnellum, Phellodon, Sarcodon. Bibliotheca Mycologica Band 104. Breitenbach, J. and Kränzlin, F. (1986). Fungi of SwitzerlandVol.2 Verlag Mykologia. Britzelmayr, M. (1910). Revision der Diagnosen zu den von Britzelmayr aufgestellen Hymenomycetenarten. P. Bech. Bot. Centralblatt. 26, 2 : 205-225. Gulden,G & Hanssen E.W.(1992). Distribution and ecology of stipitate hydnaceous fung in Norway. Sommerfeltia 13 Johannesson, H., Ryman, S., Lundmark, H., Danell, E. (1999). Sarcodon imbricatus and S. squamosus - two confused species. Mycol. Res. 103 (11): 1447-1452. Maas Geesteranus, R.A.(1975). Die terrestrischen Stachelpilze Europas (The terrestrial hydnums of Europe) Amsterdam. Pegler, D.N, Roberts, P.J & Spooner, B.M.(1997). British Chanterelles and Tooth Fungi. R B G Kew.
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A FIELD KEY TO BRITISH NON-RESUPINATE HYDNOID FUNGI Gordon Dickson Flagstones, 72 Catisfield Lane, Fareham, Hants PO15 5NS
T
he key employs characters discernible in the field. With two exceptions, all species have been examined in the fresh condition: the exceptions are Bankera violascens (of which there has been only one British record) and Hydnellum auratile. Hydnellum ferrugineum has not been seen in active growth. In the past, insufficient attention has been paid to the tree associates. The associated trees given are the ones under which I, personally, have found the species.Where there could be doubt due to the presence of more than one tree species within root range I have not drawn any conclusion.The term ‘mixed woodland’ has no meaning in this context as the fungus is mycorrhizal with only one tree species at a time. It must also be remembered that in different countries different host trees are found. This key refers only to Britain. In order to reinforce the field identification, some microscopic features have been appended. KEY TO GENERA
1) -
Growing on pine cones, stipe thin, tall, excentric . . . . . . . . . . . . . . . . . . . . .Auriscalpium vulgare Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2(1) Growing on wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 Growing on the ground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 3(2) Jelly-like, under 5 cm; dark grey to brown or fawn upper surface, pale grey spines. On conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudohydnum gelatinosum Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Hydnellum scrobiculatum with beautifully zonate and radially wrinkled caps, which may fuse together in large masses. Photograph © Gordon Dickson 99
Field Mycology Volume 1(3), July 2000 4(3)Over 5 cm diameter, white or cream, rubbery consistency; on broad-leaf trees, usually beech . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium see Key A Bracket with zoned pileus very similar in appearance to Stereum hirsutum but with buff or orange spines on underside; on hardwoods, usually beech . . . . . . . . . . . . . . . . . .Steccherinum ochraceum 5(2) White, friable, very small spines, on soil, moss or pine needles, smelling of iodoform when fresh; shape very variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sistotrema confluens Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 6(5) Cap fleshy ie. Agaricus consistency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7 Fibrous or corky consistency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 7(6) Spines more or less colour of cap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum see Key E Spines a different colour from cap surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8 8(7) Spines grey; spores brown; cap coloured. . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon see Key F Spines white to buff; spores white; cap white, then brown . . . . . . . . . . . . . . . .Bankera see Key C 9(6) Spines (and spores) white; cap black, blue-black to brown . . . . . . . . . . . . . . .Phellodon see Key D Spines (and spores) brown; cap some shade of brown +/- blue tints . . . . . . .Hydnellum see Key B Key A - HERICIUM 1 Mature spines > 3 cm. long, hanging; usually high on beech (until the tree falls!), recorded once on oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium erinaceum Mature spines < 3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 2(1) Lobed; upper surface strigose (bristly-hairy); on beech . . . . . .Hericium(= Creolophus) cirrhatum Whole fungus consisting of a mass of intertwining branches with short spines falling comb-like along their length; on beech . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hericium coralloides Key B - HYDNELLUM 1) Cap whitish, pink, buff or fulvous, darkening, consisting of fused pileoli, velvety at first; darkening with age, “lumpy” but not wrinkled or markedly zoned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Cap brown with/without blue tints; wrinkled and zoned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2(1) Cap at first fulvous or buff, velvety. Later the tomentum collapses and the surface darkens; with oak or chestnut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum spongiosipes Cap surface at first whitish to pink, “lumpy” and showing fused pileoli, then darkening; in active growth may show red guttation drops; taste mild, or mealy; with pine . . . .Hydnellum ferrugineum 3(1) Cap margin, spines and flesh all showing blue colours . . . . . . . . . . . . . . . . .Hydnellum caeruleum Cap brown (no blue), surface radially wrinkled and concentrically zoned . . . . . . . . . . . . . . . . . .4 4(3) Cap flesh thin (< 2mm), almost papery; with pine . . . . . . . . . . . . . . . . .Hydnellum scrobiculatum Not this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 5(4) Cap shades of brown, thin-fleshed, often tuberculate in the centre; with oak or chestnut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum concrescens Otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 6(5) Cap flesh thick (>3mm.); surface brown, rough and with growing edge often pink with red droplets; taste intensely hot, often after a delay; with pine . . . . . . . . . . . . . . . . . .Hydnellum peckii
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Taste not hot, cap with some orange tints; flesh thick; with pine . . . . . . . . . . . . . . . . . . . . . . . . .7
7(6) Stem orange (or partly so), flesh brown tinted orange in stem only . . . . . .Hydnellum aurantiacum Stem brown, pileus flesh brown, stem flesh dull orange with extensions through to pileus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnellum auratile NB. See species notes before recording species in couplets 2, 4 or 7. KEY C - BANKERA 1 Cap and stem whitish with slight violet tint; spines pale grey; smell sweetish when fresh, of curry when dry, with spruce (only one record to date) . . . . . . . . . . . . . . . . . . . . . . . .Bankera violascens Cap surface buff or cinnamon buff; spines and stem white, then cream; with pine, often under Calluna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bankera fuligineoalba KEY D - PHELLODON 1 Stipe black, pileus some shade of “black” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Stipe and pileus some shade of brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2(1) Cap zoned with a white edge, centre commencing brown and becoming brown-black, pileoli usually fused, not overlapping, with pine, spruce or birch . . . . . . . . . . . . . . .Phellodon melaleucus Cap with sharply defined white edge, centre with blue-black silky tomentum without shades of brown. Pileoli usually overlapping, with oak in the south, pine or spruce in Scotland. (Photo on back cover) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phellodon niger 3(1) Multiple, zoned pileoli, usually fused, with pine or spruce . . . . . . . . . . . . . . .Phellodon tomentosus Cap buff colour greying with age, not or only slightly zoned, stipe tomentose, with broad-leaf trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phellodon confluens KEY E - HYDNUM 1 Cap usually fairly robust (over 5 cm) pale cinnamon to almost cream; spines almost concolorous, decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum repandum Cap cinnamon to orange, not usually over 5cm; spines concolorous, free or adnate, NOT decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnum rufescens KEY F - SARCODON 1 Cap with recurved scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 Cap smooth, cracking in central area but without recurved scales; stem base blue-grey, both externally and in flesh; with pine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon glaucopus 2(1) Cap scaly, stem pale except at base which is blue-grey internally and externally; with sweet chestnut or oak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon scabrosus Cap with dark erect scales on a pale ground; stem pale throughout its length. With pine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sarcodon squamosus
Notes on the species in the key Some of the Hydnellum and Phellodon species are relatively easy to recognise when they are in active growth but extremely difficult when growth has ceased, even using microscopic methods. Fortunately many have an extended season and it is often possible to find a younger specimen than the old one first located. It is not worth spending time on old material. I have noted here the trees under which I have found the species. Where more than one tree has been noted within root range I have recorded this (the host tree is not always the nearest one). If the same species of tree occurs repeatedly and is never absent there is a strong probability that this is the host species. For the present I have indicated the tree which I 101
Field Mycology Volume 1(3), July 2000 am confident is the mycorrhizal partner in Britain and have shown in square brackets the species indicated by Pegler et al.(1997) for Europe. Baird(1985) quotes trees within 20m from the fungus but this is an uncertain method and I believe that it is better to use for reference those instances where there is irrefutable proof that no other species of tree is involved, even though this limits the number of observations. I would be grateful to have further information on new determinations in order to upgrade this key. In particular I would like to have notes of fungi found in pure stands of one species of tree (being careful to note the odd birch which sneaks in!) For the present it seems sufficient to note the genus of tree but if the species can be named so much the better. Auriscalpium vulgare The ‘ear-pick’ fungus cannot be confused with anything else. Always on the cones of pine (sometimes buried). Bankera fuligineoalba The combination of white stem and spines, with a variably brownish pileus surface and a ‘mushroom’ consistency leaves no doubt as to its identity. It has a very weak aroma when fresh but smells quite strongly of curry when dry; it occurs with pine and seems to prefer fruiting under the overhang of heather, especially along paths [pine]. Bankera violascens This species has been found only once (September 1996) by a track through a spruce plantation near Loch Morlich in Scotland. The spines are white but the rest of the fungus is a dull violaceous colour. It has a sweet, not unpleasant smell when fresh. The one collection was at a trackside where it had already been damaged by passing boots. It occurs only with spruce.
Hericium species On close examination the intertwined growth of H. coralloides is distinctive and the two remaining species can be separated on spine length. Even from a distance H. erinaceum can be seen either to be a round ball or with horizontal tiers whereas H. cirrhatum consists of lobes protruding in any direction. Hydnellum aurantiacum and H.auratile I have not seen fresh material of Hydnellum auratile but have seen many collections of H.aurantiacum. The latter has a stem which is orange at least in parts, there is orange on the cap surface and certainly the brown flesh is suffused with orange in the stipe. H. auratile is stated to have a brown stem, very little orange on the cap surface and the orange flesh in the stipe extends up into the pileus. Both occur only with pine. Hydnellum caeruleum Unmistakable among British species of Hydnellum; the blue coloration during growth
Hydnellum caeruleum showing the blue, tomentose margin of actively growing material. Photograph © Gordon Dickson. 102
Field Mycology Volume 1(3), July 2000 marks it out immediately. When old this colour disappears from the surface and spines but is retained in the flesh. With Caledonian pine [pine, spruce, beech]. Hydnellum concrescens and H. scrobiculatum These species are lumped together because macroscopically their appearance overlaps so much that they are almost indistinguishable. Both have radially wrinkled and concentrically zoned brown pilei which may or may not be fused. Microscopically they can be distinguished by the tubercles on the spores which in H.concrescens have a double crest (like a two-humped camel) whilst in H.scrobiculatum they are single-humped. Also the spores are slightly larger (5-6.5 x 4-5μm as against 4.5-5.5 x 3-3.5μm for H. concrescens). In the field at present I have found H. concrescens to associate with oak or chestnut and H.scrobiculatum with pine. Hydnellum ferrugineum I have not seen material of this species when in active growth, only when fully mature. It is closely related to H.spongiosipes but is a more reddish brown colour and occurs with pine. It may show red ‘guttation drops’ when actively growing but can be distinguished immediately from H. peckii which also has red drops - by the mild taste and zoned flesh [pine, spruce]. Hydnellum peckii Dark brown and thick-fleshed, this species may have a lumpy or wrinkled pileus surface but it has one distinguishing character unique in British Hydnellum species. It has an intensely hot or acrid taste, felt, after a variable delay, at the back of the throat. With experience one learns to spit out before this stage is reached! I have found it only with Caledonian pine [pine, spruce]. Hydnellum spongiosipes This species is easy to recognise: its pileus surface can best be described as ‘lumpy’ and thickfleshed. It is not wrinkled and the colour changes with the collapse of the surface tomentum from clay-buff to brown vinaceous or sepia. The concrescent pileus shows little evidence of its separate origins but the multiple stems are clad in a soft, humus - binding base. I have found it only with oak, usually in moss banks [oak, beech, chestnut]. Hydnum repandum & H.rufescens There is little difficulty in identifying Hydnum in the field but separating the two species is less easy
especially as more or less intermediate forms exist. Apart from the colour difference, note should be taken of the distribution of the spines: those in H.repandum are decurrent, often reaching half way down the stem whilst those of H.rufescens are usually free of the stem or only just reaching it. It is also a less robust fungus. Oak, beech, birch. [oak, beech, birch, maple, hazel, pine, spruce]. Phellodon confluens The distinguishing feature of this species is its unzoned pileus surface. During growth it has the usual white pileus margin of all Phellodon species but otherwise the surface is a ferruginous buff, somewhat rough and slightly wrinkled. It is relatively thick fleshed, becoming top-shaped when fully grown and with a relatively short stipe, the spines extending to the level of the moss in which it usually grows. I have found it only with oak on clay [chestnut, oak, beech]. Phellodon melaleucus & P. niger Both these species are characterised by having a sharply defined white border to the fruit-body which consists of fused pilei about 2mm. thick tapering to almost paper-thin at the edge, often giving the appearance of a single, very large cap. The underside, however, reveals multiple stems. The white border is present only during active growth. During this phase they can be distinguished in the field by the character of the cap which catches the light as it is tilted in the hand and is a blue-black colour. P. melaleucus has a duller tomentum and the colour is brown-black. In the laboratory an additional check is the partly encrusted, greening, hyphae in the context of P. niger when mounted in alkali; those of P. melaleucus are olive-green to black. I have found P. melaleucus with pine, spruce and birch on sandy soil [broadleaf and coniferous trees]. P. niger I have found under oak on heavy marls in the New Forest and under Caledonian pine and planted spruce on sandy soils in Scotland [chestnut, beech & pine]. Phellodon tomentosus Unlike P. niger in which the pileoli lose their individuality on fusing, those of P. tomentosus retain a markedly zoned appearance in shades of brown, darker at the centre and, during growth, with a white margin. They are thin and papery at the edges especially when growth has ceased. This appearance is unmistakable in the field. Under pine [pine].
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Field Mycology Volume 1(3), July 2000 Pseudohydnum gelatinosum A lumpy, tongue-shaped mass of firm, translucent jelly, grey, brown or fawn on the surface, paler underneath. The only jelly-like spine-fungus. Always on dead conifer. Sarcodon glaucopus At a casual glance this species could be mistaken for an old specimen of S.imbricatus especially as it occurs in the same habitat, but it has no clamp connections, the pileus surface is darker and not scaly (sepia/umber), the spines grey and when it is inverted the lower part of the stem is seen to be grey. This colour is also present in the adjacent flesh. It is probably not very uncommon in some areas of Scotland, having been overlooked in the past. Caledonian pine [pine]. Sarcodon scabrosus Until recently this was known as S. imbricatus in Britain, but it is now known that these are distinct species and that the latter is not British (see Field Mycology 2: 42-43). This species is the only Sarcodon occurring with broad-leaf trees in Britain. It is usually with sweet chestnut or oak in Britain (though Fries’ original description says ‘In pinetis montanis raro’). The cap surface is chestnut brown, darkening with age, the scales being adpressed except in the centre; the spines are grey and the stem base also grey but it cannot be confused with the previous species as it has adpressed or slightly recurved scales and occurs with broad-leaf trees [oak, chestnut, pine]. Sarcodon squamosus The commonest of the Sarcodon species, this occurs from Scotland to the New Forest and is invariably associated with Scots pine in Britain. It has a ‘mushroom’ consistency and the scaly cap may vary from buff to a darker cigar brown in the centre. The scales may be adpressed in the outer part of the cap to upstanding in the centre. The flesh is whitish, the spines grey and the stem white. Like other spine fungi it seems to like to fruit under the protection of heather either at tracksides or beside rivers. Microscopically it can be differentiated from S. scabrosus by having clamp connections. Other Sarcodon species One other Sarcodon has been found in Britain: Sarcodon regalis was collected (and named) from Windsor Great Park in 1968: it is now regarded as extinct in Britain. Sarcodon fuligineoviolaceus, was listed as endangered by Ing (1992), but Pegler et al. (1997) were unable to find any evidence of it
having occurred in Britain. Sarcodon leucopus has not yet been recorded in Britain. It has a smooth, pale cap and could be mistaken for a Bankera but for the coloured spines; it occurs with conifer so should be looked for in Scotland. Sistotrema confluens This is not really a spine fungus but a poroid fungus with irpicoid pores. When growing on compacted soil it sometimes makes a stem and pileus and the pores can then be seen. Growing on a bank the hymenial surface can appear hydnoid. Its fragility and smell of iodoform when fresh identify it. Known chiefly but not exclusively from pines (it has been found with oak in the New Forest) but its ecology seems uncertain. Steccherinum ochraceum A fungus frequently missed because of its resemblance to Stereum hirsutum which is so common that the underside is left uninspected. The short spines on the underside are white, cream or orange. On deciduous wood. Acknowledgements I would like to thank P.D. Orton for many helpful comments on the first draft and for supplying me with (and translating) the Britzelmayr paper referred to below and to Martin Ainsworth and Adrian Newton for further helpful comments. Literature consulted Baird, R.E.(1986). Study of the stipitate hydnums from the Southern Appalachian Mountains Genera: Bankera, Hydnellum, Phellodon, Sarcodon. Bibliotheca Mycologica Band 104. Breitenbach, J. and Kränzlin, F. (1986). Fungi of SwitzerlandVol.2 Verlag Mykologia. Britzelmayr, M. (1910). Revision der Diagnosen zu den von Britzelmayr aufgestellen Hymenomycetenarten. P. Bech. Bot. Centralblatt. 26, 2 : 205-225. Gulden,G & Hanssen E.W.(1992). Distribution and ecology of stipitate hydnaceous fung in Norway. Sommerfeltia 13 Johannesson, H., Ryman, S., Lundmark, H., Danell, E. (1999). Sarcodon imbricatus and S. squamosus - two confused species. Mycol. Res. 103 (11): 1447-1452. Maas Geesteranus, R.A.(1975). Die terrestrischen Stachelpilze Europas (The terrestrial hydnums of Europe) Amsterdam. Pegler, D.N, Roberts, P.J & Spooner, B.M.(1997). British Chanterelles and Tooth Fungi. R B G Kew.
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