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Hydnellum nemorosum A.M. Ainsw. & E. Larss. sp. nov. MycoBank MB 837985. Figs 6A, C, D, 9E. Etymology: The epithet nemorosum refers to broadleaved trees which are thought to be the mycorrhizal partners of this species. Typus: UK, Berkshire (VC22), Windsor Great Park, Johnson’s Pond (SU96676951), on a moss-covered (Leucobryum) sandy mound near Castanea sativa, with Pinus sylvestris and Quercus robur further away, 51.416389 / -0.611111, 15 Sep. 2008, A.M. Ainsworth (holotypus K(M)197478; isotypus GB-0207601); GenBank accession: MW187588. Description: Basidiomata terrestrial, stipitate, medium-sized and fleshy, solitary or clustered, ofen in small groups. Pileus 40–100 mm broad, irregularly rounded to lobate, initially convex or umbonate, becoming more plane with age and sometimes developing a depressed centre; margin thin, undulating and remaining long incurved; cuticle initially almost smooth, matt, becoming pellicular and sometimes shiny in places, occasionally showing superficial tearing to produce a few marginal areolae and zones of poorly-developed appressed scales, and more frequently becoming deeply lacerated to produce radial fissures in the underlying paler context and a central zone of coarse block-like scales; initially pinkish or vinaceous brown, with whitish growing edge, becoming progressively darker reddish or vinaceous brown towards the centre and blackening with age. Stipe 20–60 × 10–30 mm, cylindrical or basally tapered with a smooth or fibrillose texture and sometimes covered by rudimentary or entire spines; concolourous with the young pileus at the apex, darker below and distinctly bluish–green to black at the base with whitish mycelium binding the soil. Spines to some degree decurrent, up to 5 × 1 mm, pinkish brown with whitish tips at first, becoming progressively vinaceous and browner from the base. Flesh not zoned, whitish with pink tinges becoming more vinaceous when exposed to the air by tearing of pileal surface, with distinctive greyish- or bluishgreen patch within the base of the stipe, smell farinaceous but with a penetrating fruity element, taste farinaceous and bitter. Chemical reaction: when a drop of 3 % KOH is added to dry specimens, the pileipellis becomes darker brown and the flesh becomes pale brown. Hyphal system monomitic, all hyphae simple septate, tramal hyphae of spines up to 6.5 µm wide. Basidia clavate, with four sterigmata. Basidiospores brown, subglobose or short ellipsoid, irregularly tuberculate, with oblique apiculus, (3.0‒)3.4‒5.1(‒5.3) × (2.8‒)3.0‒3.8(‒4.5) µm, av. = 4.1 × 3.4 µm, Q = 1.0‒1.6 (n = 4/100, measurements from the lateral side without tubercles), tubercles numerous, 0.2‒0.8 µm high, with rounded, flat-topped or exsculpate apices. Ecology and distribution: We are of the opinion that this species is ectomycorrhizal, although we are currently unaware of any ectomycorrhizal root tip DNA evidence to confirm this.From field notes accompanying the specimens sequenced, we conclude that, although records are comparatively sparse, this species is found in mixed or broadleaved woodland and is therefore almost certainly a mycorrhizal partner of various broadleaved tree species. Hosts are likely to include Castanea sativa, Fagus sylvatica and Quercus spp. It should be noted, however, that there were no representatives of Fagaceae recorded near our species in the mixed woodland present at the sampled Swedish locality (see Additional specimens examined). Our choice of epithet for this species therefore reflects our greater degree of uncertainty regarding the range of its mycorrhizal partners compared to our state of knowledge regarding H. fagiscabrosum. It has been found in sandy or gravelly soils,sometimes with other stipitate hydnoids, and seemingly with a preference for raised or sloping ground or similarly nutrient poor microhabitats.The true extent of its European distribution is currently unknown due to the current paucity of known collections. It may have been misdetermined as a range of Sarcodon species in the past, but there are so few records to date that it could well be a genuine rarity. Thus far, we have produced molecular evidence for its presence in northern and western Europe (Denmark, Norway, Sweden and the UK), although this is currently restricted to just a single site in each country. Additional specimens examined: Denmark, Jylland, Silkeborg, Kobskov, on mineral soil on sloping ground near Fagus sylvatica, 15 Sep. 2017, T. Borgen DMS-9211777 (GB-0195939) (as S. regalis). Norway, Aust-Agder, Grimstad, Bakken Nord, on soil in steep southwest facing Quercus - dominated broadleaved woodland, 19 Sep. 2011, I.-L. Fonneland O-F-242352 (as S. ioeides). Sweden, Dalarna, Rättvik, Rättviksheden, on soil amongst mosses in mixed forest (but no Fagus or Quercus present), 8 Sep. 2000, D. Broström GB-0195631 (as S. fuligineoviolaceus). UK, same details as for holotype, 25 Sep. 2010, A.M. Ainsworth K(M)197481 (as Sarcodon sp.) Notes: This species has been described and illustrated online under the name Sarcodon regalis sensu Strandberg & Borgen in Danmarks svampeatlas (Strandberg & Borgen 2020) wherein it was documented repeatedly at a single Danish location between 2002 and 2019. It has also been recognized as “ Sarcodon sp. 2 ” in an unofficial species conservation assessment (Data Deficient) in the UK (Smith et al. 2016). Two other names (see above) have been misapplied to this species in Norway and Sweden and both are likely to have arisen due to the dominant dark purplish brown pigmentation of its mature basidiomata. We do not know if this species occurs in southern Europe but, if it is present in any historical collections, there is at least one further name that should be included in the search, namely S. cyrneus. This is another broadleaved tree associate which was described from two Corsican collections (Maas Geesteranus 1975a, b). Its protologue refers to its pinkish and purplish brown colours and poorly defined pileal scales, morphological characters which are reminiscent of H. nemorosum, but, critically, the two species differ in the pigmentation at the base of the stipe. Indeed, Maas Geesteranus (1975a) keys his species in section Squamiceps rather than section Scabrosi based on its lack of bluish or greenish colours in the stipe base. Furthermore, the illustration of the dried holotype in Maas Geesteranus (1975a) seems too pale to be H. nemorosum and there is also a difference in spore length. Bearing in mind that Maas Geesteranus’ (1971) spore measurements of Hydnellum and Sarcodon always included the ornamentation, he stated that the length of “probably not mature” spores of S. cyrneus ranges from (5.8‒)6.3‒7.3 µm whereas the corresponding values obtained for our specimens of H. nemorosum (n = 100) are 4.5‒5.8(‒6.0) µm. It would have been very interesting to include authentic S. cyrneus in our analyses but, unfortunately, our attempts to generate a sequence from type material were unsuccessful (see Discussion for further remarks on S. cyrneus).

Hydnellum roseoviolaceum Nitare sp.nov. MycoBank MB837986. Figs 7A‒C, 9F. Etymology: Epithet derived from roseo (L) = rose, referring to the rosy (rose-coloured) basidiomatal context, combined with violaceum (violet-coloured) referring to its tendency to change colour to violet-lilac. Typus: Sweden, Härjedalen, Sveg, Fisktjärnområdet, Ytterberg, in dry, lichen-dominated, seminatural old pine heath forest on acidic sand together with Cladonia spp., 62.072398 / 14.541843, 19 Sep. 2009, B. Petterson & S. Pratheepchuang (holotypus GB-0195936; isotypi O, UPS); GenBank accession: MW144374. Description. Basidiomata terrestrial,stipitate,fleshy and compact, simple, ofen rather small and slender. Pileus (30‒)50‒80 mm broad, convex to plano-convex, somewhat depressed in the centre, with undulating edge, ochraceous brown, tobacco/cigarbrown to reddish brown, with blackish spots in old or damaged parts. Pileus becoming more-or-less dirty black-spotted due to blackening of the hypodermis. Cuticle (pileipellis) dry, epidermis in young basidiomata forming a very thin brown tomentum, later becoming almost smooth, with no radially arranged fibrils, but with short and very small adnate hairy tufs from agglutination of tomentum hyphae (only seen under a lens),in patches sometimes cracked into small areoles. Dry specimens sometimes with small yellowish dots of excreted matter. Stipe 40‒60 × (5‒)10‒15 mm, concolourous with the pileus, tapering downwards and with a short greyish-blue rooting point. Spines strongly decurrent, but rarely reaching the middle of the stipe,up to 5 mm long,crowded, with age dark brown, but for a long time pallid at the tip. Flesh not zoned, first pinkish to rosy, becoming violaceous to lilac in about 20 s, at least in the central parts of the basidioma, more-or-less blackening when drying. No particular smell, taste mild. Chemical reaction. When adding 3 % KOH to dry specimens, both the pileipellis and flesh (trama) immediately change colour to charcoal black. Hyphal system monomitic,all hyphae simple septate,tramal hyphae in spines up to 10 µm wide. Basidia clavate, clampless, with 4 sterigmata. Basidiospores pale brownish, globose or subglobose, tuberculate, with oblique apiculus, 4.3‒5.1 × 3.2‒4.2 µm, av. = 4.7 × 3.6 µm, Q = 1.1‒1.5 (n = 2/48, measurements from the lateral side without tubercles), tubercles numerous, 0.5‒0.8 µm high, with prominent rounded apices. Ecology and distribution: Presumably ectomycorrhizal with Pinus sylvestris. Found in old, seminatural Pinus stands (pine heaths) on dry, acidic sandy soil with Cladonia lichens. Only known from three records (two localities) in eastern central Sweden, within the middle and northern boreal vegetation zones in areas with a rather continental climate. Additional specimens examined: Sweden, Härjedalen, Sveg, Fisktjärnområdet, Ytterberg, 29 Aug. 2008, P. Hedberg (UPS) (= first record at the type locality); Dalarna, Våmhus, Kumbelnäs, Bonäsheden, old pine heath on fossil eolian sand-dunes, 8 Sep. 2008, D. Broström GB-0195687. Notes: Among the hydnoid fungi associated with Pinus sylvestris, Hydnellum roseoviolaceum seems to be very close to, and has been mistaken for, H. fuligineoviolaceum due to its tendency to assume a lilac flesh.It differs by smaller spores,colour changes in the flesh from rose to violet (not being violet from the beginning as in H. fuligineoviolaceum) and its mild taste (N.B. some people cannot detect the very acrid taste of H. fuligineoviolaceum). It differs from H. glaucopus e.g. by its more intense rose-lilac flesh and by being associated with Pinus (not Picea) on acidic, noncalcareous sites. The pictures, and partly the description, of S. fuligineoviolaceus, by Maas Geesteranus (1975a, e.g. plate 38) seem very similar to those of this new species (collections from Greece not checked) and this may be due to a misapplication of the name. To compare the newly described species with Kalchbrenner and Fries’ original concept of Hydnum fuligineo-violaceum, the Carpathian collection from pinewoods at Olaszi (now Spišské Vlachy in Slovakia) made in Sept. 1870 and bearing Fries’ handwriting (UPS, F-173546) was studied. The collection is selected as the type by Maas Geesteranus (1975a) but he refrained from a precise designation by writing “(Holo?)Typus”. This is probably the original collection Fries refers to as v. s. (= vidi siccam; I have seen it dry) (Fries 1874). The material consists of one small basidioma divided in three slices. The flesh is typically bluish-violet-grey (also in the spines) and the spores are 5.1‒6.2 × 3.4‒4.4 µm, av. = 5.6 × 3.7 µm (n = 32, measurements from the lateral side without tubercles), tubercles 0.6‒0.8 µm high. Kalchbrenner’s collection therefore represents, without doubt, the present concept of Hydnellum fuligineoviolaceum. An ITS BLAST search in GenBank brings up two western North American sequences as the closest matches at 95–96 %. These sequences are both generated from ectomycorrhizal root-tips, one from a Pinus muricata stand in northern California (Moeller et al. 2014), the other from Pseudotsuga menziesii in British Columbia (Kranabetter et al. 2015). The sequence difference to H. roseoviolaceum consists of 15 indels and 14 gaps. We currently cannot suggest any name for this American sister taxon, if any exists. If the basidiomatal colours are like those of H. roseoviolaceum, it is likely to become misidentified as H. fuscoindicum.

Hydnellum fagiscabrosum A.M. Ainsw. & Nitare, sp. nov. MycoBank MB 837984. Figs 4A, C–F, 9A. Etymology: Epithet derived from fagi -, referring to the association with Fagales (e.g. Castanea, Fagus and Quercus) and scabrosum, referring to the morphological similarity to Hydnum scabrosum Fr. Typus: Sweden, Blekinge, Ronneby, north-eastern shore of lake Listersjön, close to a picnic area, near Fagus sylvatica, 56.320946 / 15.350357, 3 Sep. 2014, J. Nitare (holotypus GB-0195805; isotypi K(M)264450, UPS); GenBank accession: MW144294. Misapplication: Sarcodon scabrosus (at least in part) sensu European authors. Selected illustrations (all labelled as S. scabrosus): Breitenbach & Kränzlin [1986: 235 (no. 279)], Maas Geesteranus (1975a: taf. 34 abb. a), Pegler et al. (1997: fig. 77 showing micromorphology of specimen K(M)30165 sequenced by Brock et al. 2009). Description: Basidiomata terrestrial, stipitate, medium to rather large and fleshy, solitary or clustered, ofen in small groups. Pileus 50–140 mm broad, irregularly rounded to lobate, initially convex, umbonate or plane, usually developing a depressed centre at maturity;margin thin,undulating and initially incurved; cuticle radially fibrillose, showing superficial tearing to produce marginal areolae and zones of concentric scales, sometimes terminating in an upturned pointed darker tip, and deeper tearing to produce radial fissures in the underlying paler context and a central zone of coarse block-like scales; initially pinkish red brown, sometimes showing lilac or pale violaceous tints, with whitish growing edge, becoming progressively darker towards the centre and entirely chestnut brown or black brown with age. Stipe 30–100 × 10–30 mm, cylindrical or basally tapered with smooth, scaly, longitudinally fibrillose texture or covered by rudimentary or entire spines; concolourous with the pileus at the apex and distinctly bluish-green to black at the base with whitish mycelium binding the soil. Spines not, slightly or strongly decurrent,up to 10 × 1 mm, light greyish brown with whitish tips at first, becoming progressively browner from the base. Flesh not zoned, whitish, with distinctive greyish- or bluish-green patch within the base of the stipe, smell farinaceous, taste farinaceous and bitter. Chemical reaction: when a drop of 3 % KOH is added to dry specimens, the pileipellis becomes darker brown and the flesh becomes pale brown. Hyphal system monomitic,all hyphae simple septate, tramal hyphae of spines up to 8 µm wide. Basidia clavate, with four sterigmata. Basidiospores brown, subglobose or short ellipsoid, irregularly tuberculate, with oblique apiculus, 4.5‒6.3(‒6.4) × (3.5‒)3.8‒5.3(‒5.6) µm, av. = 5.4 × 4.7 µm, Q = 0.9‒1.5 (n = 4/100, measurements from the lateral side without tubercles), tubercles numerous, up to 1.3 µm high, with rounded, flat-topped or exsculpate apices. Ecology and distribution: We conclude that this species is ectomycorrhizal from the placement of GenBank sequence MF946050 which was obtained from an American ectomycorrhizal root tip of Quercus section Lobatae and identified as S. scabrosus in Rasmussen et al. (2018). From field observations we infer that H. fagiscabrosum is a mycorrhizal partner of Fagus sylvatica, Quercus spp. and Castanea sativa, including coppiced non-native Castanea in the UK. It is mostly found in the nemoral vegetation zone in sandy or gravelly soils, usually with other stipitate hydnoids, and ofen on mossy embankments, tracksides, ditchsides or in similarly nutrient poor microhabitats. The true extent of its European distribution is currently unknown due to its former inclusion within the circumscription of S. scabrosus, but we have produced molecular evidence for its presence in Norway, Sweden, UK and Italy and therefore suspect it is a very widespread, albeit relatively uncommon, member of the European Fagaceae -associated stipitate hydnoid community. Placement of GenBank sequences in Fig. 3 demonstrate that H. fagiscabrosum is also present in the southeastern USA in Florida, North Carolina and Tennessee where it has been assigned to S. scabrosus (Hughes et al. 2009, Baird et al. 2013, Rasmussen et al. 2018). Additional specimens examined: Italy, Liguria, Savona, Sassello,Badami, on soil near Castanea sativa, 2 Sep. 2010, F. Boccardo K(M)197487 (as S. regalis). Norway, Agder, Tvedestrand, N of Øynesvann, trackside in Quercus forest, 23 Aug. 2014, I.-L. Fonneland & D. Pettersen O-F-251442. Sweden, Bohuslän, Lysekil par., Vägeröds dalar, on soil with Quercus sp. and Tilia cordata, E. Larsson GB-0195727; Sotenäs and Tossene par., Hogsäm, on soil under Fagus sylvatica, R.-G. Carlsson GB-0195621, GB-0195625; Tanum par., Lindö, on soil with Quercus sp., Tilia cordata, and Corylus avellana, J. Olsson GB-0195622; Västergötland, Sätila par., Ramhultafallet, on soil with Quercus sp. and Corylus avellana, R.-G. Carlsson GB-0195621. UK, Berkshire (VC22), Windsor Crown Estate, on soil near Castanea sativa, 28 Sep. 1979, R. Phillips K(M)119189 (as S. scabrosus); Windsor Crown Estate, Buttersteep area (dry ditch), (SU9065), on soil near Castanea sativa, 2 Sep. 2005, A.M. Ainsworth K(M)197477 (as S. scabrosus); Windsor Crown Estate, Buttersteep Hill, (SU9066), on soil near Castanea sativa, 29 Sep. 2008, A.M. Ainsworth K(M)197472 (as Sarcodon sp.); East Norfolk (VC27), St Faith’s Common (TG181173), on soil near Castanea sativa, 22 Sep. 2011, A. Crotty K(M)197476 (as S. scabrosus); South Hampshire (VC11), New Forest, Brock Hill area (SU27020553), on soil near Quercus sp. and Fagus sylvatica, 9 Oct. 2011, M. Nesbitt K(M)172590 (as S. scabrosus); New Forest, Roydon Woods (SU313000), on soil, 11 Sep. 2002, A. Leonard K(M)181351 (as S. scabrosus); New Forest, Vinney Ridge (SU26290518), on soil near Quercus sp., 17 Sep. 2010, A. Lucas Hyd229 (as S.scabrosus); Surrey (VC17), Witley Common (SU92553982), on soil near Quercus sp. and Castanea sativa, 13 Sep. 2008, L. Goodwin K(M)160940 (as S. scabrosus); Woking (TQ018605), on soil near Quercus sp., Castanea sativa and Pinus sylvestris, 31 Jul. 2007, R.A. Alder K(M)162048 (as S. scabrosus); ibid., 6 Sep. 2011, R.A. Alder K(M)171979 (as S. scabrosus); West Kent (VC16), Seal Chart (TQ567557), on soil near Quercus petraea, 11 Oct. 2010, J. Pitt K(M)197490; Tudeley Woods, on soil near Castanea sativa, 24 Sep. 1994, N. Fletcher K(M)30165, (as S. scabrosus); ibid., 15 Oct. 1999, J. Weightman K(M)64653 (as S. scabrosus). Notes: Historically, our species has been included in a broad concept of Sarcodon scabrosus which has an inferred association with both Fagaceae and Pinaceae, at least in Europe. However, based on its protologue, the discussion in Maas Geesteranus & Nannfeldt (1969) and the clustering of a sequence derived from its neotype with several sequences derived from coniferassociated basidiomata and mycorrhizal root samples (see Fig. 3), Hydnellum scabrosum sensu stricto was revealed to be an ectomycorrhizal partner of Pinaceae only. More specifically, it was detected in the roots of Pinus sylvestris in Estonia (UNITE UDB008050), of Pinus densiflora (GenBank AB251833) in Japan (Lian et al. 2006) and of Pseudotsuga menziesii (GenBank KM402896) in Canada (Kranabetter et al. 2015). Furthermore, European basidiomatal sequence and collection data indicate that it is usually found on poor sandy soils with P. sylvestris and never in pure stands of Fagaceae. It should also be noted in passing that the placement of GenBank sequence AF351870 in Fig. 3 indicates that, in Oregon at least, H. scabrosum sensu stricto can also form mycorrhizal associations with epiparasitic monotropoid plant roots (Bidartondo & Bruns 2001). Hitherto, H. fagiscabrosum was recognized as “ Sarcodon sp. 1 (with Fagaceae)” in the UK (Smith et al. 2016) and as “ H. fagiscabrosum nom. prov. ” in Sweden (Nitare 2019). We have not been able to find any usable existing name for this misinterpreted species, and old names such as Hydnum amarescens Quél., a fairly pale and minutely scaly species with a non-existent type fide Maas Geesteranus (1956), are all dubious and have been applied to other species (see e.g. discussion in Maas Geesteranus & Nannfeldt 1969). Therefore, we prefer to give this widely distributed species a new name. The basidiomata of H. fagiscabrosum, H. illudens and H. scabrosum are similarly coarsely scaly when fully mature, but these species differ in their pileal pigmentation and ecological associations. In the field, H. fagiscabrosum can be distinguished from H. scabrosum by the relatively persistent, ofen broad and contrastingly whitish pileal margin of the former and its association with Fagaceae. Hydnellum scabrosum has a more concolorous pileal margin and it associates with Pinaceae. Although H. illudens and H. fagiscabrosum might be found in similar habitats in southern Europe, they differ in basidiomatal colours. The pileal surface is yellowish brown in H. illudens and the dried flesh also assumes a yellowish colour, whereas the pileus is reddish brown in H. fagiscabrosum and the flesh remains greyish when dried. Furthermore, basidiomata of H. illudens have a strongly farinaceous taste, whereas those of H. fagiscabrosum are rather sour and acrid. Hydnellum fagiscabrosum is distinguished from H. nemorosum and H. lepidum, both of which are associated with broadleaved trees, by its more coarsely scaly pileus. Of these species, H. nemorosum differs the most with its pinkish to vinaceous-brown pileus which is fissured but not really scaly, whereas in H. lepidum the scales are small and more or less adpressed all over the pileal surface., Published as part of Nitare, J., Ainsworth, A. M., Larsson, E., Parfitt, D., Suz, L. M., Svantesson, S. & Larsson, K. - H., 2021, Four new species of Hydnellum (Thelephorales, Basidiomycota) with a note on Sarcodon illudens, pp. 233-254 in Fungal Systematics and Evolution 7 (1) on pages 238-245, DOI: 10.3114/fuse.2021.07.12, http://zenodo.org/record/5838834, {"references":["Breitenbach J, Kranzlin F (1986). Fungi of Switzerland vol. 2. Verlag Mykologia, Switzerland.","Maas Geesteranus RA (1975 a). Die terrestrischen Stachelpilze Europas [The terrestrial hydnums of Europe]. 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Svensk Botanisk Tidskrif 63: 401 - 440.","Lian C, Narimatsu M, Nara K, et al. (2006). Tricholoma matsutake in a natural Pinus densiflora forest: correspondence between above- and below-ground genets, association with multiple host trees and alteration of existing ectomycorrhizal communities. New Phytologist 171: 825 - 836.","Kranabetter JM, Stoehr M, O'Neill GA (2015). Ectomycorrhizal fungal maladaptation and growth reductions associated with assisted migration of Douglas-fir. New Phytologist 206: 1135 - 1144.","Bidartondo MI, Bruns TD (2001). Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure. Molecular Ecology 10: 2285 - 2295.","Smith JH, Suz LM, Ainsworth AM (2016). Red List of Fungi for Great Britain: Bankeraceae, Cantharellaceae, Geastraceae, Hericiaceae and selected genera of Agaricaceae (Battarrea, Bovista, Lycoperdon & Tulostoma) and Fomitopsidaceae (Piptoporus). http: // fungi. myspecies. info / content / conservation.","Nitare J (2019). Skyddsvard skog. Naturvardsarter och andra kriterier for naturvardesbedomning [Woodland nature conservation - indicator species and other criteria for assessing the protection value of woodland sites]. Skogsstyrelsen, Sweden.","Maas Geesteranus RA (1956). The stipitate hydnums of the Netherlands I. Sarcodon P. Karst. Fungus 26: 44 - 60."]}

Hydnellum illudens (Maas Geest.) Nitare, comb. nov. MycoBank MB 837988. Figs 5A‒D, 9C. Basionym: Sarcodon illudens Maas Geest., Proc. K. Ned. Akad. Wet., Section C 79 (3): 285. 1976. Synonym: “ Sarcodon pseudoglaucopus ”, Nitare nom. prov. in Nitare & Högberg (2012). Misapplication: Sarcodon glaucopus sensu Nitare, 2006, non Maas Geesteranus & Nannfeldt, 1969. Selected illustrations (as “ Sarcodon pseudoglaucopus ” Nitare nom. prov.): Nitare & Högberg (2012: fig. 9A‒F). Typus: France, Vendée, Saint-Hilaire-de-Talmont, ” Le Veillon ”, under an old Quercus ilex, 4 Nov. 1973, J. Boiffard L.09111973 (holotype). Description: Basidiomata terrestrial, stipitate, fleshy and compact, single or concrescent. Pileus 70‒120 mm, convex to plano-convex, centrally somewhat depressed, ochraceous to fulvous brown, sometimes darker in the middle. Cuticle initially slightly tomentose, dry, first loosely covered by a thin cottony tomentum-layer of ephemeral whitish to pale rose hyphae (mostly whitish, but sometimes with slightly pinkish tints, ofen seen only in the expanding margin), with age becoming smooth, more or less glabrescent, from the centre more or less cracked into areoles and small adnate scales. Stipe 40‒60 × 15‒30 mm, above concolourous with the pileus, tapering downwards with a short rooting point, at the base white-tomentose or greyish blue to olivaceous grey (usually only at the rooting point, but sometimes developing a blue-grey base of the stipe). Spines strongly decurrent, but rarely reaching the middle of the stipe, up to 5 mm long, crowded, at first pallid, then becoming yellowish brown, when dried yellowish to ochraceous. Flesh not zoned, when fresh pale greyish, when dry (exsiccates) yellowish to ochraceous. Smell subfarinaceous, taste at first strongly farinaceous, mild or slightly acid or bitterish, afer a short while leaving a more intensely bitter taste with farinaceous components. Chemical reaction. When adding 3 % KOH on dry specimens, only the pileipellis (not the flesh) immediately changes colour to charcoal black. Hyphal system monomitic, all hyphae simple septate, spine trama hyphae up to 7 µm wide. Basidia clavate, with four sterigmata. Basidiospores pale brownish, globose or irregularly subglobose, tuberculate, with oblique apiculus, 4.7‒5.7(‒6.1) × 3.5‒4.5 µm, x = 5.2 × 4.0 µm, Q = 1.2‒1.5 (n = 3/90, measurements from the lateral side without tubercles), tubercles numerous, 0.4‒0.9 µm high, with rounded, flat-topped or exsculpate apices. Ecology and distribution: Associated with Picea, Pinus or Quercus on calcareous or somewhat base-rich soils, ofen in sandy and dry places. In addition to the type locality in France it is known from calcareous ground in several districts in Sweden within the hemiboreal, southern boreal and middle boreal vegetation zones. It is confirmed also from southern to central Norway, Estonia, Switzerland, and Italy. A sequence (UDB024054) deposited in the UNITE database under the name Sarcodon imbricatus proved to be this species. The sequence was generated from a basidioma collected in conifer forest in NW British Columbia, Canada, making it the first report of Hydnellum illudens from North America. Additional specimens examined: Italy, Puglia, Bari, Casamasella 14 km E of Maglia, old forest of Quercus ilex on calcareous soil, 6 Jan. 2007, E. Arnolds GB-0195786; Sardinia, Baldo, near Quercus suber, Cistus, Arbutus, 12 Nov. 2006, C.A. Hobart K(M)197492 (as S. cyrneus). Norway, Nord-Trøndelag, Steinkjer, Kalvøya, on soil with Pinus sylvestris, H. Holien, O-F-68659; Hedmark, Hamar, Furuberget nat. res., K. & E. Bendiksen O-F-76340; Viken, Ringerike, SW of Ultvedt, on soil in conifer forest, G. Gaarder O-F-242769; Ultveitvatnet nat. res., on soil in conifer forest, 25 Sep. 2010, S. Svantesson GB-0195937; Ultvedtåsen, calcareous Pinus forest, 16 Sep. 2016, T.E. Brandrud & B. Dima O-F-256728; Nordbyåsen, calcareous Pinus forest with some Picea, 20 Sep. 2016, T.E. Brandrud & B. Dima O-F-256727. Sweden, Dalarna, Rättvik, Rättviksheden, “Gropen”, sandy Pinus heath, 5 Sep. 2004, J. Nitare GB-0195825 (fig. 9E & F in Nitare & Högberg 2012; ITS1 JX999975); Kalkverket SO, lichen-rich Pinus forest, 21 Aug. 2010, E. Larsson GB-0195721; Enån nat. res., Kungshol, sandy Pinus heath, 7 Sep. 2018, E. Larsson GB-0195723, GB-0195724; Gotland, Gothem, Åminne, Tjälders, on soil in mixed conifer forest on limestone, 23 Aug. 2005, J. Nitare GB-0195803 (ITS: 1 JX999984); 26 Sep. 2011, J. Nitare GB-0195649; Gästrikland, Gävle, Limön, Oxharen, mixed conifer forest, 18 Aug. 2012, L. Andersson & T. Fasth GB-0195651; NE of the café, Picea forest with some Pinus on diabase, 18 Aug. 2012 L. Andersson & T. Fasth GB-0195808; Limön 18 Aug. 2012, L. Andersson & T. Fasth GB 0195652, GB-0195653, GB-0195654, GB-0195655; Uppland, Älvkarleby, Billudden, Brämsand, on soil in calcareous sandy Pinus sylvestris forest, 12 Sep. 2007, J. Nitare GB-0195818 (fig. 9A in Nitare & Högberg 2012; ITS: 1 JX999985); 10 Oct. 2010, G. Aronsson GB-0195650 10; 14 Sep. 2009, J. Nitare GB-0195819 (fig. 9B & 9C in Nitare & Högberg 2012); Västland, Östervret, calcareous Picea forest, 16 Sep. 2016, J. Nitare GB-0195802; Öland, Böda, Böda kronopark, Pinus forest on windblown coastal sand-dunes, 8 Oct. 2007, E. Arnolds GB-0195787. Notes: This species differs macroscopically from Hydnellum scabrosum, H. fagiscabrosum and H. glaucopus by its ochraceous to fulvous brown cuticle (almond-brown or like café-au-lait) without copper-rufous (reddish) or violaceous colours and in dried exsiccates by its yellowish-ochraceous flesh and hymenium (spines). Both H. illudens and H. scabrosum can associate with Pinus, but H. illudens is found on base-rich soils whereas H. scabrosum is found mostly on acidic soils. Hydnellum illudens usually produces smaller and more compact and sturdier basidiomata compared to H. scabrosum. Hydnellum glaucopus can also be distinguished from the others by its smaller basidiospores which are 4‒4.5 µm, evenly subglobose and ornamented with rather slender evenly spread spines that are rarely twinned. By contrast, the larger basidiospores (up to 6.5 µm long) of H. fagiscabrosum, H. illudens and H. fagiscabrosum are slightly irregular and ornamented with rather coarse, frequently twinned (exsculpate) spines. Maas Geesteranus (1976) regarded H. lepidum (as Sarcodon) as the most closely related species.This observation is confirmed by the molecular phylogeny where the two species cluster together with high support. Since its description from a single specimen the species has been little reported and seemingly ignored or misunderstood by European mycologists. The specimens we have sequenced were either not fully identified or assigned to Sarcodon cyrneus, S. fennicus, S. glaucopus, S. regalis, S. scabrosus, S. versipellis or to the provisional name “ S. pseudoglaucopus ”. Two sequenced Italian specimens(GB-0195786,K(M)197492) were collected under Quercus ilex and Q. suber, respectively, which is also the normal habitat for Sarcodon cyrneus. The possibility that H. illudens can associate with Quercus trees implies that ecology alone may not suffice to separate H. illudens from H. lepidum. See also notes to H. nemorosum below., Published as part of Nitare, J., Ainsworth, A. M., Larsson, E., Parfitt, D., Suz, L. M., Svantesson, S. & Larsson, K. - H., 2021, Four new species of Hydnellum (Thelephorales, Basidiomycota) with a note on Sarcodon illudens, pp. 233-254 in Fungal Systematics and Evolution 7 (1) on pages 245-246, DOI: 10.3114/fuse.2021.07.12, http://zenodo.org/record/5838834, {"references":["Nitare J, HOgberg N (2012). Svenska arter av falltaggsvampar (Sarcodon) - en preliminar rapport [The genus Sarcodon in Sweden - a preliminary report]. Svensk Mykologisk Tidskrif 33: 2 - 49.","Nitare J (2006). Atgardsprogram for bevarande av rodlistade fjalltaggsvampar (Sarcodon) [Action plan for red-listed species of Sarcodon in Sweden]. Naturvardsverket, Rapport 5609, Sweden.","Maas Geesteranus RA, Nannfeldt JA (1969). The genus Sarcodon in Sweden in the light of recent investigations. Svensk Botanisk Tidskrif 63: 401 - 440.","Maas Geesteranus RA (1976). Notes on hydnums. X. Proceedings van de Koninklijke Nederlandse Akademie van Wetenschappen Ser. C. 79: 273 - 289."]}

Hydnellum scabrosellum Nitare sp. nov. MycoBank MB 837987. Figs 8A‒C, 9G. Etymology: Epithet derived from scabros [um] and - ellum (L. dim.) referring to “the small scabrosum ” (= Hydnum scabrosum Fr.). Typus: Sweden, Uppland, Börstil parish, Tvärnö, Tuskö, Tuskösundet nat. res. (“Återvändan”), in herb-rich conifer forest dominated by Picea abies with some scattered old Pinus sylvestris, on strongly calcareous moraine on the edge of an old abandoned mine-shaf, 60.22267 / 18.49009, 13 Sep. 2012, J. Nitare (holotypus GB-0195689; isotypi O, UPS); GenBank accession: MW144379. Description: Basidiomata terrestrial, stipitate, thin and slender, single or concrescent. Pileus 30‒60(‒100) mm broad, planoconvex to depressed, with undulating thin margin. Cuticle fibrillose and scaly, cracking up from the centre, at first rosy to pinkish, with whitish growing edge, with age becoming coppery or vinaceous reddish-brown to purplish-brown, with darkening blackish-brown scales in the centre. Stipe 30‒60 × 5‒10 mm, above ochraceous pinkish-brown or concolourous with the pileus, tapering downwards, at the base bluish-grey or blackishgreen under a white mycelial cover. Spines strongly decurrent, ofen reaching the middle of the stipe, up to 5 mm long, crowded, at first pallid and whitish, by age becoming yellowish brown, when dry yellowish to ochraceous. Flesh not zoned,fresh pale, when dry (exsiccates) yellowish-ochraceous brown. Smell subfarinaceous, taste bitterish. Chemical reaction. When adding 3 % KOH to dry specimens, only the pileipellis (not the flesh) immediately changes colour to charcoal black. Hyphal system monomitic, all hyphae simple septate, generative hyphae up to 10 µm wide. Basidia clavate, with four sterigmata. Basidiospores pale brownish, globose or irregularly subglobose, tuberculate, with oblique apiculus, 5.1‒6.6 × 3.4‒4.7 µm, av. = 5.8 × 4.0 µm, Q = 1.2‒1.7 (n = 3/90, measurements from the lateral side without tubercles), tubercles numerous, 0.6‒0.9 µm high, with prominent, rounded, flattened, flat-topped to exsculpate apices. Ecology and distribution: Presumably ectomycorrhizal with conifers (mixed forest with Pinus and Picea, associated treespecies not exactly known but probably Pinus sylvestris), strongly calciphilous. Mostly known from coniferous forests situated close to the coast (Gulf of Bothnia) in the province of Gästrikland and northern Uppland, east-central Sweden (at the northern limits of the hemiboreal vegetation zone). This so far Swedish species can be expected to occur in pine forests on calcareous ground around the Baltic Sea. Additional specimens examined: Sweden, Gästrikland, Hamrånge, Bergby, Näset nat. res., calcareous conifer forest, 15 Sep. 2011, J. Nitare GB-0195807; Uppland, Börstil, Tvärnö, Tuskö, Tuskösundet,5 Sep. 2008, J. Nitare GB-0195806 (from the same spot as the holotype, fig. 12A in Nitare & Högberg 2012); Älvkarleby,Lanforsen, Tippön,calcareous Pinus sylvestris forest (with scattered Picea abies), 20 Sep. 2013, G. Aronsson GB-0195791; Askön, calcareous Pinus sylvestris forest (with scattered Picea abies), 13 Sep. 2013, G. Aronsson GB-0195792; Östergötland, Västra Tollstad, Omberg, Ombergsliden nat. res., on soil in calcareous Picea abies forest with Pinus sylvestris and Betula sp., 13 Sep. 2016, E. Larsson GB-0195736. Notes: Among the hydnoid fungi associated with conifers, Hydnellum scabrosellum is most similar to H. scabrosum but has smaller and slenderer basidiomata and looks like a dwarf form of the latter species. The size difference is particularly obvious when considering the stipe diameter which rarely exceeds 1 cm in H. scabrosellum but is usually 2‒5 cm in mature specimens of H. scabrosum. The yellowish-ochraceous hymenium and flesh (when dry) and the occurrence on base-rich soil, also differs from the non-calciphilous H. scabrosum. The coastal region in Gästrikland, where most collections were made, is characterized by its extremely calcareous-rich soils (moraine), depending on the geological features and recent land uplif during the postglacial period.