Heterotypic synonym: Inocybe reducta J.E. Lange, Fl. Agaric. Danic. 5(Taxon. Consp.): V, 101 (1940).

Selected description and iconography: Velenovský (1920–1922); Lange (1940, as “I. reducta”); Kuyper (1986); Stangl (1989).

Typification

Lectotype designated by Kuyper (1985: 376): CZECH REPUBLIC, Prag, Mnichovice, in Betuletum above railway (viam ferream), Aug. 1915 [leg. J. Velenovský] (PRC-bottle n°119).

Epitype (designated here; MBT 10014037) NETHERLANDS, Drenthe, Assen, park of hospital, alt. 15 m, Quercus robur, 3 Oct. 2019, leg. D. Bandini & E. Arnolds (STU SMNS-STU-F-0901739; dupl. priv. herb. DB DB3-10-19-14).

Description

Pileus 20–45 mm wide, (sub)globose or subcampanulate when young, later broadly convex or expanded, often somewhat deformed, with or without a low, large umbo, margin at first slightly to strongly inflexed, later decurved and with age straight to even uplifted, and then depressed around the centre; no remnants of a velipellis observed; colour mostly brown with reddish to red-brown hue (Mu 7.5YR 4/4–4/6, 5/4–5/8), rarely somewhat paler and with ochraceous tinge (Mu 10YR 5/4–5/8), sometimes darker to almost blackish brown at the centre and often somewhat speckled with fine darker fibre bundles on paler ones, margin sometimes much paler with age; surface at first glabrous, then finely innately fibrillose to rimulose near the margin, but often also strongly rimose with fibres diverging and showing the lighter-coloured context beneath; when aging at the centre, usually areolate-diffracted or somewhat cracked or “scabby” or breaking up in small particles; no cortina observed. Lamellae moderately crowded [c. 40–50 (60), l = 1–3], often uneven, slightly thick, adnexed to adnate, usually very flat, even to subventricose, at first whitish, later beige to pale ochraceous; edge whitish. Stipe 30–60 × 2–6 mm, straight or curved and often widening towards the base, base usually thickened to (sub)bulbous, sometimes with a rather thick bulb (up to 10 mm), when young entirely covered with thin whitish tomentum, later longitudinally striate or glabrous, flesh-coloured to brown or reddish brown, up to dark reddish brown or red-brown with age; pruinose along entire length, but more densely so in the upper half of the stipe and sometimes seemingly only near the apex. Context whitish in the pileus, at first whitish in the stipe, with age brownish in the cortex and sometimes streakily brownish also in the stipe, base remaining whitish. Smell spermatic, at least when cut. Colour of exsiccata. Pileus brown to dark brown with reddish hue (Mu 5YR 4/4–4/6, 3/3–3/4), lamellae somewhat lighter, stipe concolorous or a little lighter, no darkening or blackening on drying.

Spores 5.9–9.5 µm (av. 7.3 µm, SD 0.7 µm) × 3.7–5.0 µm (av. 4.3 µm, SD 0.3 µm); Q = 1.4–2.1 (av. 1.7, SD 0.1) (n = 120 of 3 coll.), smooth, subamygdaloid, sometimes oblong, occasionally with a faint suprahilar depression, apex mostly subobtuse. Basidia 25–30 × 7–9 µm, generally 4-spored. Pleurocystidia 36–79 µm (av. 56 µm, SD 10 µm) × 11–24 µm (av. 15 µm, SD 3 µm); Q = 2.3–6.1 (av. 3.7, SD 1.0) (n = 45 of 3 coll.), mostly subfusiform to (sub)utriform, usually without or with only a short neck, at the apex generally wide, with a short pedicel, apex usually crystalliferous, mostly rather thin-walled and walls rather uniformly thick near bulge and apex, but with only few small crystals, walls up to 1.5 (2.0) µm thick at the apex, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size, intermixed with colourless (sub)clavate, thin-walled paracystidia. Pileipellis constituted by an epicutis made up of parallel hyphae 4–11 µm wide, with finely encrusting and parietal brown pigment, subcutis with wider and paler to colourless elements. Caulocystidia descending the entire length of the stipe but sometimes sparsely so on the lower half, 30–65 × 9–20 µm, mostly ventricose (sub)utriform or (sub)fusiform, sometimes deformed, without or with only a short neck and with a short pedicel, apex with or without small crystals; walls up to 0.5 (1.0) µm thick at the apex, pale yellowish-greenish with 3% KOH; intermixed with colourless, (sub)clavate, thin-walled cauloparacystidia. Clamp connections abundant in all tissues.

Habitat and distribution

Inocybe albomarginata was found by us with frondose trees, often in park-like terrain on calcareous soil. Judging from the collection data, the species appears to often associate with Quercus. Our own collections are from Austria, Germany and The Netherlands. The corresponding species hypothesis (SH0740134.09FU, 2.0 %) includes sequences from Czechia, Estonia, Finland, Germany, Italy, Latvia, The Netherlands, Pakistan, Russia, Sweden and Turkey. The species seems to be widespread, ranging from Europe to south Asia and quite common.

Type specimens examined

Lectotype of I. albomarginata (PRC-bottle n°119) (Fig. 8a): Spores 6.4–7.8 µm (av. 7.1 µm, SD 0.3 µm) × 3.8–4.5 µm (av. 4.2 µm, SD 0.2 µm); Q = 1.6–1.9 (av. 1.7, SD 0.1) (n = 40), smooth, subamygdaloid, sometimes oblong, occasionally with a faint suprahilar depression, apex subobtuse. Basidia 4-spored. Pleurocystidia 32–58 µm (av. 46 µm, SD 7 µm) × 11–17 µm (av. 14 µm, SD 2 µm); Q = 2.5–5.2 (av. 3.3, SD 0.6) (n = 40)], mostly (sub)fusiform or (sub)utriform, apex usually crystalliferous, walls up to 1.0 (1.5) µm thick at the apex, yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size. Paracystidia (sub)clavate, colourless, thin-walled. Caulocystidia not studied (because in a bad condition).

Syntype of I. reducta (Fig. 8c): DENMARK, Husmandskolen near Odense, Sep. 1938, [leg. J.E. LANGE] (C-F-67157). Spores 6.5–8.0 µm (av. 7.2 µm, SD 0.3 µm) × 4.0–5.0 µm (av. 4.3 µm, SD 0.2 µm); Q = 1.5–1.8 (av. 1.6, SD 0.1) (n = 40), smooth, subamygdaloid, sometimes oblong, occasionally with a faint suprahilar depression, apex subobtuse. Basidia 4-spored. Pleurocystidia 44–57 µm (av. 51 µm, SD 5 µm) × 13–17 µm (av. 15 µm, SD 1 µm); Q = 2.9–3.8 (av. 3.4, SD 0.3) (n = 40), mostly (sub)fusiform or (sub)utriform, apex usually crystalliferous, walls up to 1.0 (1.5) µm thick at the apex, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size. Paracystidia (sub)clavate, colourless, thin-walled. Caulocystidia not studied (to preserve the material).

Epitype of I. albomarginata (Figs. 2a, d): Spores 6.2–9.1 µm (av. 7.2 µm, SD 0.5 µm) × 3.7–4.8 µm (av. 4.3 µm, SD 0.3 µm); Q = 1.4–1.9 (av. 1.6, SD 0.1) (n = 40), smooth, subamygdaloid, sometimes oblong, occasionally with a faint suprahilar depression, apex (sub)obtuse. Basidia 4-spored. Pleurocystidia 42–71 µm (av. 54 µm, SD 10 µm) × 13–17 µm (av. 14 µm, SD 1 µm); Q = 2.5–5.5 (av. 3.8, SD 0.9) (n = 40), mostly (sub)fusiform or (sub) utriform, apex usually crystalliferous, walls up to 1.5 (2.0) µm thick at the apex, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size. Paracystidia (sub) clavate, colourless, thin-walled. Caulocystidia on entire length of the stipe, 35–60 × 9–15 (20) µm, mostly ventricose (sub)utriform or (sub)fusiform.

Selected additional specimens examined

AUSTRIA, Tirol, Imst, Silz, ÖK25V 2221-Ost, alt. 660 m, riparian forest with Tilia sp., Pinus sylvestris, Fraxinus sp., Populus sp., 11 Sep. 2017, leg. D. Bandini (SMNS-STU-F-0901740, DB11-9-17-13). GERMANY, Baden-Württemberg, Karlsruhe, Erzbergerstr., TK25 6916/3, alt. 118 m, lawn with Quercus petraea, Fagus sylvatica, Carpinus sylvaticus, 1 Jun. 2013, leg. D. Bandini & B. Oertel (DB1-6-13-19). Nordrhein-Westfalen, Siegen-Wittgenstein, Freudenberg, Rest stop Siegerland, TK25 5013/3, Fagus sylvatica, Corylus avellana, 21 Oct. 2013, leg. D. Bandini (DB21-10-13-1). NETHERLANDS, Drenthe, Assen, park of hospital, alt. 15 m, Quercus robur, 3 Oct. 2019, leg. D. Bandini & E. Arnolds (DB3-10-19-14). Drenthe, Gijsselte, alt. 15 m, Quercus robur, 6 Oct. 2020, leg. D. Bandini (DB6-10-20-6). Drenthe, Diever, alt. 16 m, Pinus sylvestris, Picea abies, Quercus robur, Fagus sylvatica, 8 Oct. 2020, leg. D. Bandini (DB8-10-20-1). Ibidem, alt. 15 m, Pinus sylvestris, Picea abies, Quercus robur, Fagus sylvatica, 8 Oct. 2020, leg. D. Bandini (DB8-10-20-13).

Notes

The lectotype of I. albomarginata is preserved in formaldehyde in a bottle together with samples of different genera (see also Kuyper 1985). The consistency of the fruiting bodies is soft and jelly-like and the lectotype is not in a good condition, so, e.g., the caulocystidia cannot be examined. There is no chance that caulocystidia will be observable on any other material in the bottle. Therefore, and because no drawing or detailed description nor a photograph are given in the original description, and because I. albomarginata is often mistaken for I. amblyospora, we decided to select an epitype for I. albomarginata.

What is termed above as “syntype” of I. reducta is the only type material with details matching the holotype we are aware of, but as LANGE did not specify where the type was deposited and not all of his material is in C, we cannot be sure it is the only instance of the material. Inocybe reducta was already synonymized with I. albomarginata by Kuyper (1985, 1986). Our morphological study of the type material in C supports this. Both have the same on average very small and especially very narrow spores, which excludes a synonymy with either I. amblyospora (see below) or I. pseudoreducta [with wider, often (sub)ovoid, apple-seed-like spores], and the shape of the hymenial cystidia is very similar; molecular data are not available from the type of I. reducta, because it was preserved in formaldehyde for some time.

Macroscopically, the pileus of I. albomarginata is mostly of reddish brown to red-brown colour, with a darker to almost blackish brown centre, its surface is at most finely innately fibrillose to rimulose, the lamellae are flat, and the species is furthermore characterized by an entirely but in the lower half sometimes sparsely pruinose stipe with a thickened to bulbous base and, on average, rather short subfusiform to (sub)utriform hymenial cystidia with a wide neck. A velipellis is lacking. Inocybe amblyospora, with which I. albomarginata has been mistaken [among others by Bandini et al. (2019b; MH366616) and Ryberg et al. (2008; AM882908.2), differs, e.g., by the presence of a velipellis, smoother pileus centre, on average longer spores and a different ecology. It occurs in montane to subalpine regions, often in somewhat moist terrain. Inocybe suecica Vauras & E. Larss., another species with small spores, can be distinguished from I. albomarginata, e.g., by the presence of a velipellis, smoother pileus surface and glabrous centre of pileus, as well as larger spores (Vauras & Larsson 2016; DB, personal observation), and I. pseudoreducta, e.g., by the presence of a velipellis, smoother centre of pileus, and wider spores often shaped like apple seeds (Stangl & Glowinski 1981; La Rosa et al. 2017). Inocybe bonii (see below) differs from I. albomarginata, e.g., by the greyish velipellis, smoother pileus surface at the centre, not flat lamellae and larger spores. Inocybe demetris Bandini & U. Eberh. differs, e.g., by the abundant velipellis, paler pileus colour, smooth centre of pileus, on average larger spores and much longer caulocystidia, as well as growth with conifers (Bandini et al. 2022b). Inocybe metrodii Stangl & J. Veselský can be distinguished from I. albomarginata, e.g., by the often larger basidiomata, presence of a velipellis, smoother centre of pileus, lamellae and stipe often partly becoming yellowish when bruised, and on average larger spores (Stangl & Veselský 1979; Bandini et al. 2021c), and I. glabrescens Velen., e.g., by the generally less stout basidiomata, abundant velipellis, pileus colour without reddish tinge, and larger spores (Velenovský, 1920-1922; Bandini et al. 2021c). Apart from I. amblyospora and I. pseudoreducta, none of the named species is particularly closely related to I. albomarginata (Fig. 1).

Two undescribed taxa are most similar to I. albomarginata in their ITS (SH1144361.09FU, in Fig. 1 represented by KR673688 and LC623536; and SH1144362.09FU, in Fig. 1 represented by KF041394). The two Asian sequences form the sister clade to I. albomarginata, which is phylogenetically well supported (98/100/1) in Fig. 1. The described species most similar in the ITS, but with less than 90% similarity rather distant, are I. flavoalbida Matheny & Bougher and I. praecox Kropp, Matheny & Nanagy. Inocybe flavoalbida is a small-spored species with entirely pruinose stipe. It differs from I. albomarginata, e.g., by the paler pileus colour and habitat, since it was found in mixed subtropical rainforest in Australia and Papua New Guinea (Matheny & Bougher 2017). Neither the ITS of the holotype nor that of the paratypes of I. praecox has been published, but the sequences indicated in Fig. 1 as I. praecox are identical in their LSU to the holotype of I. praecox. Inocybe praecox can be distinguished by larger and stouter basidiomata, a partially appressed squamulose pileus surface and clavate to fusiform-ventricose hymenial cystidia. It is a vernal species and was described from western Washington state (USA); the sequences used here for the species stem from across the border, from British Columbia (Canada).

Description and iconography: Kühner (1955).

Typification

Lectotype designated by Poirier (2016: 54): FRANCE, Savoie, above Avenches, “Pierre Larron” [alt. 1770 m], moist location with Picea abies, under Adenostyles, 18 Aug. 1937 [leg. R. Kühner] (G00058748).

Description

Pileus 20–40 mm wide, at first (sub)conical to subcampanulate, later conico-convex, convex or expanded, with or without a more or less pronounced large umbo, margin at first involute or decurved, later decurved to straight or even uplifted and then pileus depressed around the centre; when young, with faint remnants of a greyish velipellis, visible sometimes also later at the centre of the pileus; colour dark brown with reddish hue to dark reddish brown (Mu 5YR 4/3–4/6, 3/3–3/4; 7.5YR 3/4), sometimes up to blackish brown at the centre; surface at first smooth and shiny glabrous, later still shiny and smooth or slightly verrucose at the centre, outwards minutely appressed floccose or rim(ul)ose to minutely innately fibrillose; young basidiomata with remnants of a cortina. Lamellae from moderately crowded to subdistant (c. 40–50, l = 1–3), broadly adnate, subventricose, at first whitish to whitish-greyish, later pale yellowish to pale ochraceous-brownish; edge strongly fimbriate, whitish. Stipe 30–60 × 3–7 mm, rather stout, cylindrical or widening towards the base, base thickened to (sub)bulbous, at first entirely covered by fine whitish tomentum, later longitudinally striate to glabrous, at first very pale brownish, then brownish to intensely red-brown, near the base and at the extreme apex remaining whitish and thus sometimes with strong colour contrast; rather roughly pruinose over its entire length. Context whitish in pileus and stipe, brownish in the cortex of the stipe when old. Smell indistinct. Colour of exsiccata. Pileus brown to dark brown with reddish hue (Mu YR 7/5YR 4/4–4/6, 3/4), lamellae a little lighter in colour, stipe concolorous or a little darker in colour, no darkening or blackening on drying.

Spores 7.2–10.0 µm (av. 8.4 µm, SD 0.4 µm) × 3.9–5.0 µm (av. 4.4 µm, SD 0.2 µm); Q = 1.6–2.3 (av. 1.9, SD 0.1) (n = 120 of 3 coll.), smooth, mostly narrow (sub)ellipsoid or subcylindrical, with or without a suprahilar depression, apex obtuse, rarely subobtuse. Basidia 25–30 × 7–9 µm, generally 4-spored, rarely also 2-spored and then spores up to 10 µm. Pleurocystidia 39–74 µm (av. 58 µm, SD 8 µm) × 12–23 µm (av. 17 µm, SD 3 µm), Q = 1.7–5.6, av. 3.6, SD 0.9) (n = 45 of 3 coll.), mostly (sub)utriform, also (sub)fusiform or subcylindrical, with sometimes a rather long neck, usually with a short, but sometimes with a long pedicel, apex usually with only small crystals, walls up to 1.0 (1.5) µm thick at the apex, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size, intermixed with numerous colourless (sub)clavate, (sub)cylindrical or subglobose, thin-walled paracystidia. Pileipellis constituted of an epicutis made up of parallel hyphae 5–12 µm wide, with encrusting and parietal brownish to brown pigment, subcutis with wider and paler to colourless elements, intermixed with thin-walled colourless cauloparacystidia. Caulocystidia on the entire length of the stipe, 35–60 × 10–15 (20) µm, broadly (sub)fusiform, (sub)utriform, rarely (sub)clavate, apex usually crystalliferous, walls up to 1.0 µm thick at the apex, pale yellowish-greenish with 3% KOH, intermixed with colourless (sub)clavate, thin-walled cauloparacystidia. Clamp connections abundant in all tissues.

Habitat and distribution

Inocybe amblyospora was found by us with Picea abies at high montane to subalpine altitudes, often in rather moist conditions. One collection was found on a small island in the middle of a brook, another on a moist slope crisscrossed by springlets. This fits very well to what Kühner wrote about the location of the holotype (see below). Besides our own collections from Austria and Germany, no other sequences of basidiomata or EcM or soil-samples are known to us. Thus, the species seems to be rather rare.

Type specimens examined

Lectotype (G00058748). Spores 7.3–10.0 µm (av. 8.8 µm, SD 0.7 µm) × 3.6–4.8 µm (av. 4.4 µm, SD 0.3 µm); Q = 1.7–2.4 (av. 2.0, SD 0.2) (n = 40), smooth, oblong, mostly (sub)cylindrical, apex obtuse. Basidia 4-spored. Pleurocystidia 52–75 µm (av. 62 µm, SD 8.0 µm) × 8–19 µm (av. 14 µm, SD 3.1 µm); Q = 3.1–6.5 (av. 4.7, SD 1.0) (n = 15), (sub)cylindrical, (sub)fusiform or subutriform, apex usually crystalliferous, walls up to 1.0 (1.5) µm thick, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size. Paracystidia not observed. Caulocystidia not studied (stipe was not sent by the herbarium).

Additional specimens examined

AUSTRIA, Tirol, Imst, Silz, Kühtai, ÖK25V 2221-Ost, alt. 1780 m, Picea abies, 20 Aug. 2012, leg. D. Bandini & B. Oertel (DB20-8-12-12). Salzburg, Tamsweg, Prebersee, ÖK25V 3230-Ost, alt. 1500 m, moist location with Picea abies, Alnus sp., 10 Aug. 2013, leg. D. Bandini (SMNS-STU-F-0901742, DB10-8-13-11). Salzburg, Zederhaus, Riedingtal, ÖK25V 3229-Ost, alt. 1490 m, moist location with Picea abies, 18 Sep. 2020, leg. D. Bandini (DB18-9-20-25). GERMANY, Bayern, Ostallgäu, Pfronten, Breitenberg, TK25 8429/3, alt. c. 1800 m, very moist location with Alnus viridis, Picea abies, 19 Aug. 2012, leg. D. Bandini & B. Oertel (DB19-8-12-15). Bayern, Ostallgäu, Halblech, Ammergebirge, TK25 8431/1/3, alt. 1400 m, Picea abies, 17 Sep. 2018, leg. D. Bandini (SMNS-STU-F-0901741, DB17-9-18-9).

Notes

Inocybe amblyospora is characterized by a dark brown to dark reddish brown pileus colour, smooth to rim(ul)ose or minutely fibrillose pileus surface, faint greyish velipellis, stipe pruinose over entire length, smooth, on average short and narrow spores, mostly (sub)utriform hymenial cystidia often with a rather long neck, and broadly (sub)fusiform or (sub)utriform caulocystidia. It can be found at montane to subalpine altitudes with conifers. The species has been mistaken for I. albomarginata (see under I. albomarginata), which differs from I. amblyospora, e.g., by lacking a velipellis, by the often areolate-diffracted or somewhat cracked pileus centre, flat lamellae and shorter spores, as well as a habitat usually at lower altitudes (see above). Inocybe bonii has, e.g., on average shorter but wider spores, on average smaller hymenial cystidia and longer, mostly narrow utriform or subcylindrical caulocystidia with mostly a long and undate neck. The spores and hymenial cystidia of I. suecica are on average wider, and the species grows with frondose trees (Vauras & Larsson 2016; DB, personal observation). Inocybe brijunica Mešić, Tkalčec & Haelew. can be distinguished from I. amblyospora, e.g., by stipe pruinose only near the apex or at least not in the lower half, much shorter spores and shorter hymenial cystidia, as well as a Mediterranean habitat (Mešić et al. 2021), and I. demetris, e.g., by the abundant velipellis, on average wider spores, hymenial cystidia often with undate walls, and much longer caulocystidia (Bandini et al. 2022b). Inocybe pseudoreducta differs, e.g., by pileus surface more decidedly rimose with age, spores often shaped like apple-seeds with (sub)conical apex, as well as habitat usually at lower altitudes (Stangl & Glowinski 1981; La Rosa et al. 2017).

The two available sequences for I. amblyospora received high ufb and pp support (100and 1, respectively) in Fig. 1. The species is clearly distinct from all other described or unnamed taxa. Inocybe amblyospora is, with around 90% similarity in the ITS, only very distantly related to I. praecox, which has larger basidiomata than I. amblyospora, a paler pileus colour, on average wider spores, clavate to ventricose fusiform hymenial cystidia, and occurs in spring (Kropp et al. 2010 and see above under I. albomarginata). We were not allowed to sequence the lectotype of I. amblyospora, but since the macroscopic and microscopic details of our own collections match well with those of the type, and since the habitat is the same, we are confident that the collections we sequenced are I. amblyospora and consider our sequences suitable as surrogates for a type sequence, at least until data from the type become available.

Etymology

Named “bonii” in honour of Marcel Bon, who was an outstanding inocybologist.

Diagnosis

Inocybe bonii has a brown pileus colour, mostly with reddish hue, at most rimose pileus surface, entirely pruinose stipe, smooth spores, size on average 8.3 × 5.0 µm, mostly (sub)fusiform hymenial cystidia, pleurocystidia measuring on average 52 × 13 µm, and narrow utriform or subcylindrical caulocystidia with mostly a long and undate neck. It grows on calcareous soil. It can be recognized by the combination of the above characteristics and differs in its ITS sequence from other species, such as I. albomarginata or I. pseudoreducta.

Holotype

GERMANY, Baden-Württemberg, Alb-Donau-Kreis, Merklingen, Widderstall, TK25 7424/3, alt. 850 m, Picea abies, Fagus sylvatica, 30 Sep. 2021, leg. D. Bandini (STU SMNS-STU-F-0901743; isotype priv. herb. D.B. DB30-9-21-3).

Description

Pileus 10–30 mm wide, at first (sub)conical to subcampanulate, later broadly convex or expanded, often without, otherwise with a more or less pronounced large umbo, margin at first decurved, later straight to even uplifted and then depressed around the centre; young basidiomata with remnants of a pale greyish velipellis visible even later at the centre but vanishing when moist; colour brown in different shades, mostly with reddish hue (Mu 10YR 5/4–5/6, 4/4–4/6; 7.5YR 4/4–4/6; 5YR 4/4–4/6), with age sometimes strongly paling, either towards the margin or entirely; surface at first smooth and glabrous, with age either remaining glabrous or getting from minutely rimulose to strongly rimose towards the margin, with diverging fibres so that the paler context below is visible; no remnants of a cortina observed. Lamellae moderately crowded (c. 35–60, l = 1–3), adnate, sometimes with subdecurrent tooth, subventricose, long time whitish, later greyish-whitish and with age greyish-brownish with a more or less intense ochraceous hue; edge often irregular, even, whitish. Stipe 30–70 × 2–5 mm, straight or curved, base even to thickened or even submarginate bulbous, when young entirely covered with thin whitish tomentum, later longitudinally striate to glabrous, at first whitish, then flesh-coloured to pale reddish or even rather intense reddish-brownish, base/bulb whitish; pruinose on entire length, though sparsely in the lower half. Context whitish in the pileus, faintly brownish or reddish-brownish in the cortex of the stipe, whitish in the base/bulb. Smell (sub)spermatic, at least when cut. Colour of exsiccata. Pileus brown to dark reddish brown (Mu 5YR 4/3–4/6, 3/2–3/4; 7.5YR 4/4–4/6), lamellae and stipe concolorous, no darkening or blackening on drying.

Spores 7.3–9.2 µm (av. 8.3 µm, SD 0.4 µm) × 4.5–5.6 µm (av. 5.0 µm, SD 0.2 µm); Q = 1.4–2.0 (av. 1.6, SD 0.1) (n = 120 of 3 coll.), smooth, (sub)amygdaloid, mostly without a suprahilar depression, apex (sub)acute, with indistinct pseudoporus. Basidia 21–28 × 7–10 µm, generally 4-spored, rarely also 2-spored and then spores up to 10.5 µm. Pleurocystidia 41–65 µm (av. 52 µm, SD 6 µm) × 9–18 µm (av. 13 µm, SD 2 µm); Q = 2.8–6.7 (av. 4.2, SD 0.9) (n = 45 of 3 coll.), mostly (sub)fusiform, sometimes also (sub)utriform or sublageniform, usually without or with only short neck but occasionally also with a longer neck, walls sometimes somewhat undate, mostly with a rather short pedicel, apex usually crystalliferous, walls up to 2.5 (3.0) µm thick at the apex, yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size, intermixed with numerous colourless, (sub)clavate, thin-walled paracystidia. Pileipellis constituted of an epicutis made up of parallel hyphae 4–11 µm wide, with encrusting and parietal brown to dark brown pigment, subcutis with wider and paler to colourless elements, up to 15 µm wide. Caulocystidia on entire length of the stipe, but only sparsely in the lower half, 50–95 × 8–13 µm, narrow utriform or subcylindrical, mostly with a long neck with undate walls, with a short pedicel or sometimes with truncate base, apex usually crystalliferous, walls up to 1.5 µm thick at the apex, yellowish-greenish with 3% KOH, intermixed with (sub)clavate to (sub)cylindrical, thin-walled cauloparacystidia. Clamp-connections abundant in all tissues.

Habitat and distribution

Inocybe bonii was found by us on calcareous soil with conifers or frondose trees, in France and Germany. In GenBank and UNITE (SH0570962.09FU, 2.0% = SH0017908.09FU, 3%) there are no other sequences from basidiomata, but EcM sequences from Germany (HF675192), Switzerland (MN970792), and soil-samples from Estonia, Greece, Iran, Israel, Italy and Morocco that might also represent I. bonii.

Additional specimens examined

FRANCE, Charente-Maritime, Lozay, bois d'Essouverts, 46.028158 N, -0.537226 W, alt. 88 m, Pseudotsuga menziesii, Picea abies, 20 Nov. 2019, leg. F. VALADE (DB20-11-19-Valade, FV2019112002). GERMANY, Sachsen-Anhalt, Börde, Weferlingen, TK25 3632/3, Quercus robur, Carpinus betulus, 14 Oct. 2013, leg. E. Preikschas (SMNS-STU-F-0901747, DB14-10-13-Preikschas). Bayern, Ostallgäu, Füssen, cemetery, TK25 8430/1, alt. 800 m, Picea abies, 22 Sep. 2021, leg. D. Bandini (SMNS-STU-F-0901744, DB22-9-21-8). Baden-Württemberg, Alb-Donau-Kreis, Merklingen, Widderstall, TK25 7424/3, alt. 850 m, Picea abies, Fagus sylvatica, 30 Sep. 2021, leg. D. Bandini (SMNS-STU-F-0901745, DB30-9-21-5).

Notes

Inocybe bonii is characterized by an often reddish-hued brown, smooth to rimose pileus surface, pale greyish velipellis, when aging an up to intensely reddish-brownish stipe pruinose on its entire length but only sparsely so in the lower half, on average rather small smooth spores and mostly (sub)fusiform hymenial cystidia as well as narrow utriform or subcylindrical caulocystidia with a mostly long and undate neck. The species can be mistaken for several other species with small spores such as I. albomarginata, which differs, e.g., by lacking a velipellis, by the often areolate-diffracted or somewhat cracked pileus centre, flat, long time whitish lamellae and smaller spores (see above). Inocybe amblyospora differs, e.g., by the usually darker pileus colour, when aging less rimose pileus surface, the reduced and very fugacious velipellis, and narrower, oblong, often (sub)cylindrical spores with (sub)obtuse apex as well as habitat usually in higher montane areas (see above), and I. brijunica, e.g., by the usually darker brown pileus colour without reddish tinges, smaller spores and Mediterranean habitat (Mešić et al. 2021). Inocybe comis Bandini & B. Oertel has a finely tomentose to minutely or strongly lanose pileus, stipe pruinose only near the apex, spores mostly with an explicit suprahilar depression and caulocystidia often with subcapitate apex (Bandini et al. 2022b), whereas I. demetris usually has fewer reddish tinges and never reddish pilei, mostly (sub)utriform to (sub)lageniform hymenial cystidia sometimes with undate neck, on average longer spores and longer hymenial cystidia (Bandini et al. 2022b). Inocybe glabripes Ricken differs from I. bonii by usually lacking reddish tinges in the pileus colour, at most a finely rimose pileus surface, stipe pruinose only near the apex, smaller spores and shorter hymenial cystidia (see, e.g., Ricken 1910–1915; Lange 1917, as “I. microspora”; Kuyper 1986; Stangl 1989; Bandini et al. 2021c), I. minimispora, e.g., by the only faint reddish hue in the pileus colour, fugacious whitish velipellis, stipe pruinose only near the apex, smaller spores and shorter caulocystidia (Reumaux 1986; Bandini et al. 2021c), and I. heterosemen, e.g., by the at the centre reddish and towards the margin yellow-ochraceous pileus colour, on average smaller spores and shorter, often (sub)clavate hymenial cystidia (Carteret & Reumaux 2012; Bandini et al. 2022b). Inocybe pseudoreducta has on average shorter, mostly ovoid or apple-seed shaped spores and plumper, on average wider hymenial cystidia and mostly (sub)fusiform, often somewhat deformed caulocystidia not with an undate neck, whereas I. suecica never has an almost reddish pileus colour, pileus often much darker at the centre than towards the margin, stipe not becoming intensely reddish-brownish with age but paler, and hymenial cystidia often with a clearly demarcated transition from the bulge to the neck, and usually growing on more rich and less dry terrain (Vauras & Larsson 2016; DB, personal observation). The pileus of I. virgatula Kühner has at most a faint reddish hue, the stipe is pruinose only near the apex, has larger spores with a suprahilar depression and on average larger hymenial cystidia (Kühner 1955; Stangl 1974; Bandini et al. 2021c).

Based on available data, it is not clear whether soil sequences included in the same sh as sequences that quite clearly represent I. bonii all represent the species or whether I. bonii has a very close unnamed sister taxon. The distribution data could suggest that I. bonii s. str. has a temperate distribution and some of the sequenced soil samples a more Mediterranean one, but this is based on a very small number of samples and could be misleading. We here decided to treat I. bonii in the narrower sense. Phylogenetically, based on ITS and LSU, the closest relative is I. demetris (see Fig. 1); in terms of ITS similarity, I. comis and I. glabripes rank almost the same as I. demetris (all around 93%).

Etymology

Named “carolina” (“belonging to Carolus”, i.e., Carl) after Carl von Linné.

Diagnosis

Inocybe carolina has a brown to dark brown pileus colour, long time smooth and glabrous, with age only minutely innately fibrillose pileus surface, abundant whitish velipellis, stipe pruinose only near the apex, smooth spores measuring on average 9.2 × 5.6 µm, a usually rather plump and rather thin-walled, variously shaped hymenial cystidia, and pleurocystidia measuring on average 68 × 14 µm. It grows on dry calcareous soil and is probably associated with Pinus. It can be recognized by the combination of the above characteristics and differs in its ITS sequence from other species, such as I. venerabilis or I. rufobrunnea.

Holotype

SPAIN, Comunidad de Madrid, Guadalix de la Sierra, 40.774700 /-3.640960, alt. 827 m, in an abandoned limestone quarry, at mossy location close to Pinus sylvestris, Populus nigra and Juniperus oxycedrus, 26 Nov. 2018, leg. A. Díaz-Fernández, J.A. Rodea-Butragueño, M. García-Aranda & F.J. Rodríguez-Campo (holotype MA-Fungi 98710; isotypes priv. herb. F.J. RodrÍguez Campo PRC-181126-05, STU SMNS-STU-F-0901789, priv. herb. D.B. DB26-11-18-Rodr.-Campo).

Description

Pileus 5–35 mm wide, at first subconical to subcampanulate or subglobose, later broadly convex or expanded, usually without an umbo, margin at first decurved, later straight or even uplifted when old, and then pileus depressed around the umbo; young basidiomata thickly covered by a layer of whitish velipellis, visible also later at least at the centre of the pileus; colour brownish to dark brown (Mu 10YR 4/3–4/6, 3/4–3/6; 7.5 YR 3/4), but mostly somewhat speckled because of remnants of the velipellis; surface at first and often long time smooth and glabrous, later sometimes minutely innately fibrillose with fine fibres; young basidiomata with faint remnants of a whitish cortina. Lamellae moderately crowded (c. 35–50, l = 1–3), broadly adnate with a (sub)decurrent tooth, even to (sub)ventricose, at first whitish, later creamy to greyish-whitish, with age greyish-brownish; edge strongly fimbriate, whitish. Stipe 20–40 × 2–7 mm, often rather stout, straight to curved, sometimes widening towards the base or base widened or even bulbous, when young covered with whitish tomentum, later longitudinally striate to glabrous, at first pale flesh-coloured, later ochraceous brownish, with or without reddish tinge, and pale to sometimes rather intensely reddish near the apex; roughly pruinose only near the apex of the stipe. Context watery whitish in the pileus and at first also in the stipe, later greyish-whitish to reddish brown, at least in the cortex of the upper half of the stipe. Smell indistinct or subspermatic to slightly disagreeable. Colour of exsiccata. Pileus dark brown (Mu 7/5YR 3–4; 10YR 3/3–3/6), lamellae and stipe concolorous or a little lighter in colour, no darkening or blackening on drying.

Spores 8.0–10.6 µm (av. 9.2 µm, SD 0.6 µm) × 4.8–6.8 µm (av. 5.6 µm, SD 0.4 µm); Q = 1.4–2.0 (av. 1.7, SD 0.1) (n = 160 of 4 coll.), smooth, often rather heterogeneous in size, mostly broadly (sub)amygdaloid, usually without and only rarely with a faint suprahilar depression, apex mostly (sub)acute. Basidia 24–42 × 8–13 µm, generally 4-spored, rarely also 2-spored. Pleurocystidia 45–91 µm (av. 68 µm, SD 8 µm) × 10–19 µm (av. 14 µm, SD 2 µm); Q = 3.1–7.2 (av. 4.9, SD 1.0) (n = 60 of 4 coll.), rather plump (sub)utriform, (sub)cylindrical, subfusiform or (sub)clavate, without or with only a short neck, often with more or less undate walls, sometimes with a rounded and occasionally cap-like thickened apex, with a short pedicel or with truncate base, apex crystalliferous or not, walls up to 1.0 (1.5) µm thick at the apex, but mostly rather thin-walled, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in size but somewhat more variable in shape, intermixed with numerous colourless, (sub)clavate, thin-walled, sometimes catenate paracystidia. Pileipellis constituted of an epicutis made up of parallel hyphae 4–12 µm wide, with encrusting and parietal dark brown pigment, subcutis with wider and paler to colourless elements. Caulocystidia only near the apex of the stipe, 35–70 × 10–15 µm, often rather plump (sub)fusiform, (sub)cylindrical to (sub)clavate, apex with or without small crystals, walls up to 1.0 µm thick at the apex, pale yellowish-greenish with 3% KOH; intermixed with some colourless, (sub)clavate, thin-walled cauloparacystidia. Clamp-connections abundant in all tissues.

Habitat and distribution

Inocybe carolina was found by us in Austria and Spain, always on dry calcareous soil. In Austria it was found on the gravelly border of a small road, not far from the river Lech, in the vicinity of Helianthemum, Dryas and various species of gentians and orchids. The collections from Spain were found in an abandoned limestone quarry. This calcareous area is a small hill located next to the Pedrezuela reservoir, with Quercus ilex as the predominant tree species and various other species at the foot of the quarry: Populus nigra, Juniperus oxycedrus and Pinus sylvestris. Species hypothesis SH1235752.09FU, 1.5% presumably represents I. carolina, with a specimen sequence from Italy (JF908222) and soil sample sequences from Morocco. Probably not all sequences in the corresponding species hypothesis at 3% (SH0200216.09FU) represent I. carolina.

Additional specimens examined

AUSTRIA, Tirol, Reutte, Rieden, ÖK25V 2215-West, alt. 790 m, Pinus sylvestris, Juniperus sp., 8 Sep. 2019, leg. D. Bandini (DB8-9-19-22). Tirol, Reutte, Weißenbach am Lech, at the wayside, ÖK25V 2215-West, alt. 870 m, Pinus sylvestris, 9 Oct. 2021, leg. D. Bandini (DB9-10-21-7). SPAIN, Comunidad de Madrid, Guadalix de la Sierra, 40.77769 /-3.64081, alt. 830 m, in an abandoned limestone quarry, mossy location near Pinus sylvestris, Populus nigra and Juniperus oxycedrus, 16 Nov. 2015, leg. J. A. Rodea-Butragueño & A. Díaz-Fernández (MA-Fungi 98709, priv. herb. PRC-151116-02, SMNS-STU-F-0901790, DB16-11-15-Rodr.-Campo). Ibidem, 23 Oct. 2020, leg. J.A. Rodea-Butragueño & F. J. Rodríguez-Campo (MA-Fungi 98711, priv. herb. PRC-201023-01, DB23-10-20-Rodr.Campo). Ibidem, 27 Oct. 2021, leg. J.A. Rodea-Butragueño, A. Díaz-Fernández & F.J. Rodríguez-Campo (MA-Fungi 98712, priv. herb. PRC-211027-01, DB27-10-21-Rodr.-Campo).

Notes

Inocybe carolina is characterized by the brown to dark brown pileus colour, long time smooth and glabrous, with age only minutely innately fibrillose pileus surface, abundant whitish velipellis, stipe pruinose only near the apex, smooth broadly (sub)amygdaloid spores and usually rather plump and rather thin-walled, differently-shaped hymenial cystidia, as well as rather plump (sub)fusiform to (sub)clavate caulocystidia. The habitat was for all our collections dry calcareous soil with Pinus nearby. In Austria, I. venerabilis Bandini, B. Oertel & U. Eberh. was found in the same habitat. This species differs from I. carolina, e.g., by the smaller basidiomata, greyish velipellis, larger spores and often shorter hymenial cystidia (Bandini et al. 2022b). Inocybe aurantiobrunnea Esteve-Rav. & García Blanco, a species described from Spain, differs, e.g., by the less abundant velipellis, orange to orange-yellow lamellae and larger spores (Esteve-Raventós et al. 2003; Bandini et al. 2022a), while I. costinitii Bizio, Ferisin & Dovana, also a Mediterranean species, has larger spores, often subcylindrical thicker-walled hymenial cystidia and usually long and narrow (sub)cylindrical to (sub)fusiform caulocystidia (Bizio et al. 2016; Bandini et al. 2021c). Inocybe rufobrunnea J. Favre can be distinguished from I. carolina, e.g., by the more fragile basidiomata, less smooth pileus surface, greyish velipellis and much larger spores (Favre 1955; Kuyper 1986, as “I. rufuloides var. exilis”; Bandini et al. 2022b), I. tarda Kühner, e.g., by the sometimes almost blackish brown pileus colour, less abundant and darker velipellis, larger spores and longer, often somewhat deformed caulocystidia often with undate walls (Kühner 1955; Bandini et al. 2021c, 2022a), I. filiana, e.g., by the usually smaller and more fragile basidiomata, with age often less smooth pileus surface, less abundant velipellis and shorter and not plump hymenial cystidia (Bandini et al. 2022b). Furthermore, it can be found on more acidic soil. Inocybe aerea E. Ludw. has more fragile basidiomata, a less smooth pileus surface, fugacious and less abundant greyish velipellis and narrower spores, and it grows near Salix (Ludwig 2017; DB, personal observation), whereas, I. furfurea Kühner differs from I. carolina, e.g., by the usually paler pileus colour, with age scabby, cracked and/or fissured pileus centre, fugacious and less abundant greyish velipellis and on average much shorter spores (e.g., Kühner 1955; Kuyper 1986; Stangl 1989; Bandini et al. 2019c). Inocybe griseovelata Kühner usually has a paler pileus colour, more greyish velipellis, longer spores and very long and narrow caulocystidia (Kühner 1955; Kuyper 1986; Stangl 1989; Bandini et al. 2021c, 2022a), and I. grusiana Bandini & B. Oertel can be distinguished from I. carolina, e.g., by the greyish velipellis, longer spores and long and narrow, subcylindrical caulocystidia (Bandini et al. 2021c).

Inocybe aurantiobrunnea is the most similar described species to I. carolina, with a 94% sequence identity in the ITS. As for I. bonii, without morphology it is difficult to delimit I. carolina clearly against EcM and soil data, although in this case the support for I. carolina s. str. is, with 89/100/1, better than for the clades including soil or EcM sequences. In Fig. 1, I. carolina is interpreted in the strict sense.

Etymology

Named “centesima” because it is the hundredth new species described by D. Bandini and co-authors.

Diagnosis

Inocybe centesima has an at the centre pale brownish, outwards dark brown pileus colour with a purple hue, smooth to at most rim(ul)ose pileus surface, entirely pruinose stipe, smooth, mostly almost kidney-shaped spores measuring on average 11.6 × 6.0 µm, mostly (sub)fusiform hymenial cystidia, pleurocystidia measuring on average 55 × 19 µm, and growing on sandy calcareous soil with Pinus sylvestris. It can be recognized by the above combination of characteristics and differs in its ITS sequence from other species, such as I. psammobrunnea or I. rufobrunnea, that can be found in the same habitat.

Holotype

GERMANY, Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, near Nature Reserve Düne Pferdstrieb, TK25 6617/4, alt. 113 m, Pinus sylvestris, 25 Oct. 2017, leg. D. Bandini (STU SMNS-STU-F-0901749; isotype priv. herb. D.B. DB25-10-17-2).

Description

Pileus 20–35 mm wide, at first (sub)conical, later broadly convex or expanded, with or without low, large umbo, margin slightly decurved, later straight or even uplifted, and then pileus depressed around the umbo; young basidiomata with faint and fugacious remnants of a pale greyish velipellis; usually somewhat bicoloured, pale ochraceous brownish to pale nut-brown at the centre and sometimes with age also at the margin, outside the centre brown with a more or less intense purplish hue (Mu 10YR 5/6–5/8; 5YR 5/6–5/8; 10R 3/4–3/6); surface at first smooth and glabrous, later finely rim(ul)ose towards the margin; no remnants of a cortina observed. Lamellae moderately crowded (c. 40–60, l = 1–3), irregular, adnate, (sub)ventricose, at first whitish then dingy whitish with greyish hue or greyish to greyish-brownish; edge fimbriate, whitish. Stipe 40–60 × 3–5 mm, cylindrical or widening towards the base, base always thickly covered with sand, equal or slightly thickened; at first covered with fine whitish tomentum, later longitudinally striate or glabrous, at first whitish, later dingy whitish to ivory or beige; pruinose on entire length. Context dingy or watery whitish in the pileus, whitish in the stipe. Smell spermatic, at least when cut. Colour of exsiccata. Pileus dark brown with reddish hue (Mu 5YR 3/3–3/4; 7.5YR 3/4), lamellae a little lighter and stipe much paler in colour, no darkening or blackening on drying.

Spores 10.0–13.6 µm (av. 11.6 µm, SD 0.8 µm) × 5.3–7.1 µm (av. 6.0 µm, SD 0.3 µm); Q = 1.7–2.3 (av. 1.9, SD 0.1) (n = 120 of 3 coll.), smooth, mostly very characteristically almost kidney-shaped, with a suprahilar depression, apex subacute to subobtuse, with small indistinct pseudoporus. Basidia 25–30 × 7–11 µm, generally 4-spored, but also 2-spored. Pleurocystidia 37–73 µm (av. 55 µm, SD 8 µm) × 14–24 µm (av. 19 µm, SD 2 µm); Q = 1.9–4.9 (av. 2.9, SD 0.6) (n = 45 of 3 coll.), usually ventricose (sub)fusiform, at the apex generally wide, without or with only a short neck, usually without or with a short pedicel, apex usually crystalliferous, walls up to 1.5 (2.0) µm thick at the apex, pale yellowish-greenish with 3% KOH. Cheilocystidia similar in appearance and size, intermixed with numerous colourless, (sub)clavate or subcylindrical, thin-walled paracystidia. Pileipellis constituted of an epicutis made up of parallel hyphae 4–12 µm wide, with encrusting and parietal brownish to dark brown pigment, subcutis with wider and paler to colourless elements. Caulocystidia on the entire length of the stipe, but only sparsely in the lower half due to abrasion through sand particles, 30–55 × 10–18 µm, mostly (sub)lageniform or (sub)fusiform, apex usually crystalliferous, without or with only a short neck and pedicel, walls up to 1.5 µm thick at the apex, pale yellowish-greenish with 3% KOH; intermixed with numerous (sub)clavate to subglobose cauloparacystidia. Clamp connections abundant in all tissues.

Habitat and distribution

Inocybe centesima was found by us on calcareous sandy terrain (inland dunes) with Pinus sylvestris in Germany. No other collections or sequences in databases are known to us.

Additional specimens examined

GERMANY, Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, Nature Reserve Galgenbuckel, TK25 6617/4, alt. 112 m, Pinus sylvestris, Helianthemum nummularium, 31 Oct. 2021, leg. D. & G. Bandini (SMNS-STU-F-0901750, DB31-10-21-1). Baden-Württemberg, Rhein-Neckar-Kreis, Walldorf, TK25 6617/4, alt. 120 m, Pinus sylvestris, 6 Nov. 2021, leg. D. & G. Bandini (SMNS-STU-F-0901787, DB6-11-21-2). Baden-Württemberg, Rhein-Neckar-Kreis, St. Ilgen, TK25 6617/4, alt. 115 m, Pinus sylvestris, 14 Nov. 2021, leg. D. & G. Bandini (SMNS-STU-F-0901751, DB14-11-21-1).

Notes

Inocybe centesima is characterized by an at the centre pale brownish, outwards dark brown pileus colour with purple hue, smooth to at most rim(ul)ose pileus surface, only faint and fugacious pale greyish velipellis, an entirely pruinose stipe, rather large, smooth, mostly almost kidney-shaped spores and mostly (sub)fusiform hymenial cystidia. The shape of the spores of I. centesima is very unusual and the contrasting pileus colour of older basidiomata with purple tinges is quite extraordinary as well. Two species can be found at the same location: I. psammobrunnea Bon, differing from I. centesima, e.g., by the often large greyish patch at the centre of the pileus, with age up to intensely reddish-pinkish stipe, and shorter (sub)amygdaloid and not almost kidney-shaped spores (Bon 1990; Poirier 2002, as “I. griseotarda”; Bandini et al. 2021c), and I. tarda, also growing in the same location and which can be distinguished from I. centesima by, e.g., stipe with age often entirely reddish of different intensities, on average shorter (sub)amygdaloid and not almost kidney-shaped spores, on average longer hymenial cystidia and longer caulocystidia (Kühner 1955; Bandini et al. 2021c, 2022a). Inocybe tjallingiorum Kuyper usually has a darker centre of the pileus, no purple tinges in the pileus colour, stipe with age intense brown to red-brown or even darker, and much smaller, (sub)amygdaloid to (sub)ellipsoid spores (Kuyper 1986; Stangl 1989; Bandini et al. 2021c), whereas I. serotina has a larger, paler patch of velipellis at the pileus centre, larger, mostly (sub)amygdaloid and not almost kidney-shaped spores and mostly (sub)clavate or subglobose hymenial cystidia. Furthermore, I. serotina is usually found on sea-shores (Peck 1904; Lange 1940; Kuyper 1986; DB, personal observation). The Mediterranean species I. neorufula Esteve-Rav., Macau & Ferville differs from I. centesima, e.g., by the whitish velipellis, more reddish-tinged, much smaller, not almost kidney-shaped spores and on average longer hymenial cystidia (Esteve-Raventós et al. 2012; DB, personal observation), and I. rufuloides, e.g., by the lack of purple tinges in the pileus colour, less smooth pileus surface, (sub)amygdaloid or almost elliptic, not almost kidney-shaped spores and often subcapitate on average shorter hymenial cystidia (see below). Inocybe metrodii can be distinguished, e.g., by the darker centre of pileus and lack of purple tinges in the pileus colour, much smaller, not almost kidney-shaped spores, and habitat not on inland dunes but on richer soil (Stangl & Veselský 1979; Bandini et al. 2021c), I. phaeoleuca Kühner, e.g., by the lack of purple tinges in the pileus colour and the smaller and (sub)amygdaloid, not almost kidney-shaped spores (Kühner 1955; Kuyper 1986, as “I. splendens var. phaeoleuca”; Bandini et al. 2019c), and I. splendens R. Heim, e.g., by the shorter navicular and not almost kidney-shaped spores and on average longer hymenial cystidia, as well as growth with frondose trees (Heim 1931; Kuyper 1986, as “I. splendens var. splendens”; Bandini et al. 2019c).

The species clade of I. centesima is well supported (94/99/1). We are not aware of any ITS sequences more similar to I. centesima. No species genetically closely related to I. centesima are known as yet, and the sequences forming the sister clade of I. centesima are at most 95% similar. With only 86% identity in the ITS, I. pararubens Carteret & Reumaux is the most closely, yet quite distantly related described species. It differs from I. centesima, e.g., by the darker centre of pileus and lack of purple tinges in the pileus colour, smaller, (sub)amygdaloid to (sub)navicular spores and growth with frondose trees (Carteret & Reumaux 2012; DB, personal observation). Inocybe centesima is by all indications a rare species.

Selected description and iconography: Bon (1984, 1990); Kuyper (1986); Lantieri (2005); Bougher & Matheny (2011).

Typification

Holotype: FRANCE, Somme, Cayeux-sur-Mer, Brighton-La Mollière, Pinus, 18 May 1983, leg. M. Bon, J. Vast & G. Claus, (LIP-MB83038).

Epitype (designated here; MBT 10014038): GERMANY, Baden-Württemberg, Rhein-Neckar-Kreis, Walldorf, TK25 6617/4, alt. 120 m, Pinus sylvestris, 6 Nov. 2021, leg. D. & G. Bandini (STU SMNS-STU-F-0901753; dupl. priv. herb. D.B. DB6-11-21-4).

Description

Pileus 10–40 mm wide, at first (sub)conical, later broadly convex or expanded, with or without a large umbo, margin at first slightly inflexed, soon deflexed, later straight or even uplifted when old and then depressed around the umbo; young basidiomata entirely covered by a greyish velipellis, still visible at the umbo of older basidiomata; colour brown to dark brown, often with reddish hue in different nuances (Mu 10YR 4/3–4/6, 3/4–3/6; 5YR 4/3–4/6; 7.5YR 4/4–4/6), but when young often greyish-brownish because of the velipellis; surface at first sometimes almost smooth, but usually finely tomentose to minutely lanose, with age coarsely lanose, sometimes somewhat lacerate; young basidiomata with faint remnants of a cortina. Lamellae rather distant (c. 25–35 (40), l = 1–3), thickish, narrowly to broadly adnate, (sub)ventricose, at first whitish, later greyish-brownish to brown with reddish hue; edge uneven fimbriate, whitish. Stipe 15–45 × 2–5 mm, stout, straight or curved, base often somewhat thickened, at first entirely covered with whitish tomentum, later longitudinally striate or glabrous, reddish brown at different intensities to intensely red-brown, base mostly remaining whitish; pruinose with sparse, rough pruina only near the apex of the stipe. Context whitish in the pileus, red-brown in the upper half of the stipe. Smell spermatic, at least when cut. Colour of exsiccata. Pileus pale brown, greyish-brownish (Mu 10YR 5/4–5/6, 4/2–4/6), lamellae and stipe concolorous or a little lighter in colour, no darkening or blackening on drying.

Spores 9.7–13.4 µm (av. 11.4 µm, SD 0.8 µm) × 5.5–7.1 µm (av. 6.4 µm, SD 0.3 µm); Q = 1.4–2.1 (av. 1.8, SD 0.1) (n = 120 of 3 coll.), smooth, oblong (sub)amygdaloid or almost elliptic, with or without a faint suprahilar depression, apex subacute to subobtuse, with indistinct pseudoporus. Basidia 25–32 × 9–12 µm, generally 4-spored, rarely also 2-spored. Pleurocystidia 34–69 µm (av. 48 µm, SD 7.3 µm) × 10–20 µm (av. 14 µm, SD 2.1 µm); Q = 2.3–4.8 (av. 3.4, SD 0.6) (n = 45 of 3 coll.), variously shaped, often subcapitate, (sub)cylindrical, (sub)utriform, (sub)fusiform, (sub)lageniform), sometimes even (sub)clavate, at the apex generally wide, mostly with only a short neck, with a short pedicel or with truncate base, apex usually crystalliferous, walls up to 1.5 (2.0) µm thick at the apex, often abruptly thickened at the neck, yellow-green with 3% KOH. Cheilocystidia similar in appearance and size, intermixed with numerous colourless to sometimes somewhat brownish, (sub)clavate, (sub)cylindrical or subglobose or somewhat deformed, thin- to slightly thick-walled, sometimes catenate paracystidia. Pileipellis constituted of an epicutis made up of parallel hyphae 4–12 µm wide, with encrusting and parietal brownish pigment, subcutis with wider and paler to colourless elements. Caulocystidia only near the apex of the stipe, 30–60 × 10–15 µm, (sub)utriform, (sub)cylindrical or even (sub)clavate, but mostly somewhat deformed and often with undate walls, without or with only a short neck, without or with a short pedicel or with truncate base, apex crystalliferous or not, walls up to 1.0 (1.5) µm thick at the apex, yellow-green with 3% KOH; intermixed with numerous, mostly but sometimes somewhat brownish, (sub)clavate to subglobose, thin-walled colourless cauloparacystidia. Clamp connections abundant in all tissues.

Habitat and distribution

Inocybe rufuloides was found by us only in the calcareous sand of inland dunes or adjacent areas, always with Pinus sylvestris nearby. However, only Quercus was recorded for the collection from Algeria. The taxon appears to correspond to SH0146902.09FU, 3%, which includes specimen sequences from Algeria, Australia, Italy, Germany and Spain, an EcM sequence from France and soil sample sequences from Italy and Morocco. Based on this, the species appears to be widely distributed and not rare. Bougher & Matheny (2011) considered I. rufuloides as an introduced taxon in Australia.

Additional specimens e„xamined

ALGERIA, Bainem, Quercus ilex, 31 Dec. 2014, leg. S. Aouali (DB31-12-14-Aouali). GERMANY, Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, Nature Reserve “Düne Pferdstrieb”, TK25 6617/4, alt. 100 m, Pinus sylvestris, Quercus robur, Helianthemum, 13 Oct. 2012, leg. D. BANDINI, B. Oertel & W. Winterhoff (SMNS-STU-F-0901442, DB13-10-12-4). Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, TK25 6617/4, alt. 113 m, Pinus sylvestris, Helianthemum, Quercus robur, 21 Oct. 2012, leg. D. Bandini & B. Oertel (DB21-10-12-14). Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, Nature Reserve “Düne Pferdstrieb”, TK25 6617/4, alt. 113 m, Pinus sylvestris, 23 Oct. 2021, leg. D. & G. Bandini (DB23-10-21-2). Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, Nature Reserve “Galgenbuckel”, TK25 6617/4, alt. 112 m, Pinus sylvestris, Helianthemum nummularium, 31 Oct. 2021, leg. D. & G. BANDINI (SMNS-STU-F-0901752, DB31-10-21-5). Baden-Württemberg, Rhein-Neckar-Kreis, Sandhausen, Nature Reserve “Galgenbuckel”, TK 6617/4, alt. 112 m, Pinus sylvestris, Helianthemum nummularium, 6 Nov. 2021, leg. D. & G. Bandini (DB6-11-21-4). Baden-Württemberg, Rhein-Neckar-Kreis, Walldorf, TK25 6617/4, alt. 120 m, Pinus sylvestris, 6 Nov. 2021, leg. D. & G. Bandini (DB6-11-21-5).

Notes

A description of the holotype was given in Bandini et al. (2020c). We have tried and failed to generate PCR products from this collection; others appear to have failed, too. According to our experience, it is only rarely possible to successfully sequence material collected by M. BON. We decided to choose an epitype for the species as the type material did not display all morphological details necessary for its delimitation, for instance the caulocystidia at the top of the stipe and the paracystidia of young basidiomes. In the protologue, Bon (1984) described the latter as abundant and sometimes catenate. A plate of the microscopic details and measurements of the spores and pleurocystidia of the type were published in Bandini et al. (2020c).

Inocybe rufuloides is characterized by a usually rather stout habit, brown to dark brown pileus colour with more or less intense reddish hue, with age mostly lanose pileus surface, abundant pale greyish velipellis, stipe with age up to intensely red-brown, pruinose only near the apex, rather large spores, and often subcapitate hymenial cystidia with many, sometimes catenate and slightly thick-walled paracystidia. Inocybe carneicaulis E. Ludw. differs, e.g., by the smoother pileus surface, usually quite uniformly (sub)fusiform, usually not (sub)capitate hymenial cystidia and not catenate or slightly thick-walled paracystidia (Ludwig 2017; DB, personal observation), I. coriacea, e.g., by the smaller and less stout basidiomata, usually paler pileus colour, on average smaller spores and usually not subcapitate hymenial cystidia (Bandini et al. 2022c), and I. deianae Eyssart., e.g., by the velipellis usually not remaining at the centre of the pileus of older basidiomata, on average shorter spores and usually not subcapitate hymenial cystidia (Eyssartier 2007; Brugaletta et al. 2019, as “I. lapidicola”; Bandini et al. 2022a). Inocybe distantifolia E. Ludw. can be distinguished from I. rufuloides, e.g., by the smaller pileus, smoother pileus surface, on average smaller spores and usually not subcapitate hymenial cystidia (Ludwig 2017), and I. laurina Bandini, B. Oertel & C. Hahn, a species found by us next to I. rufuloides, can be distinguished from the latter, e.g., by the whitish velipellis, smoother pileus surface, smaller spores and longer, usually not subcapitate hymenial cystidia (Bandini et al. 2020a). Inocybe psammobrunnea has a smoother pileus surface, smaller spores and on average longer and usually not subcapitate hymenial cystidia (Bon 1990; Poirier 2002 (as “I. griseotarda”); Bandini et al. 2021c), I. neorufula has a whitish velipellis, smoother pileus surface and smaller spores (Esteve-Raventós et al. 2012; DB, pers. obs.) and I. rufobrunnea has a smoother pileus surface, larger spores and usually not subcapitate hymenial cystidia (Favre 1955; Kuyper 1986, as “I. rufuloides var. exilis”; Bandini et al. 2022b). Inocybe tarda differs from I. rufuloides, e.g., by having less velipellis, a smoother pileus surface, on average shorter spores and on average longer, usually not subcapitate hymenial cystidia (Kühner 1955; Bandini et al. 2021c, 2022a).

Inocybe rufuloides is well supported (97/100/1) and well delimited in Fig. 1, in a clade including a number of unnamed sequences. Among the described and sequenced species, I. filiana is most similar in the ITS to I. rufuloides (87%). This species differs from I. rufuloides, e.g., by the usually smaller and more fragile basidiomata, usually smoother pileus surface, smaller spores and not often subcapitate hymenial cystidia (Bandini et al. 2022b).

Basionym: Agaricus geophyllus Bull., Herb. France 11: pl. 522, fig. 2 (1791).

Typification

Lectotype, designated here (MBT 10003226): Bulliard (1791), Herbier de la France 11: pl. 522, fig. 2 (Agaricus geophyllus).

Epitype, selected in Bandini et al. (2021c: 1058): AUSTRIA, Tirol, Reutte, Höfen, ÖK25V 2215-West, alt. 970 m, Picea abies, 23 Sep. 2015, leg. D. Bandini (STU SMNS-STU-F-0901531; dupl. priv. herb. D.B. DB23-9-15-31).

Notes

This species was discussed in detail and epitypified in Bandini et al. (2021c). The here designated lectotype was cited as the holotype in the latter publication. As mentioned in the introduction, epitypes only serve as interpretative types for the material that is cited in the epitypification and not for syntypes. For purposes of epitypification, illustrations are better considered lectotypes, because illustrations cannot represent the entire type that ever existed. Furthermore, the designation of an illustration as lectotype, rather than citing it merely as holotype, protects the epitypification against other illustrations, variations of the same illustration, etc. that might exist and could be considered syntypes based on the original description. Lectotypification in cases such as this (even if no competing syntype is known at the time) avoids potential ambiguity and ensures nomenclatural stability. With this in mind, we decided to follow up the earlier epitypification with a lectotypification.

Typification

Lectotype, designated here (MBT 10014039): FRANCE, Paris, Bois de Vincennes, in a circle of more than 20 fruiting bodies on soil, with frondose trees, near the lake, 8 Jun.1930, leg. R. KÜHNER (G00058738)

Notes

The collection date of this collection agrees with the protologue. This lectotypification supersedes the lectotypification of Bandini et al. (2021c) [Art. 9.12 (Turland et al. 2018)], who selected a collection with a (presumed) earlier collection date because the above collection seemed to have been lost, when ordered by us in 2018. In the above publication, we cited a wrong G accession number for the earlier collection; the correct number for that specimen is G00566267. The collection G00058738, from June 8th 1930, was suggested earlier as lectotype by J. Poirier (herbarium note, 16 Jan. 1995), but this was, to the best of our knowledge, never formally published. With the correction of the lectotypification, also the epitype SMNS-STU-F-0901568 designated in Bandini et al. (2022a) loses its standing (Art. 9.20. Note 8). We have not had the opportunity to study the lectotype G00058738. Therefore, it would be premature to assign an epitype. The species was discussed in detail in Bandini et al. (2021c, 2022a).

Typification

Lectotype, designated here (MBT 10006582): ITALY, northern Italy. Bresadola (1884) Fungi Trident., 1(4-5): plate 58, fig. 1.

Epitype, selected in Bandini et al. (2022b: 118): GERMANY, Niedersachsen, Emsland, Haselünne, TK25 3310/4, alt. 20 m, Corylus avellana, Quercus robur, Pinus sylvestris, 4 Oct. 2020, D. Bandini (STU SMNS-STU-F-0901607; dupl. priv. herb. D.B. DB4-10-20-19).

Notes

The species was discussed in detail in Bandini et al. (2022b). The selected illustration was cited as holotype there. See under I. geophylla for an explanation of the lectotypification.

Typification

Lectotype, designated by Poirier (2016: 55): FRANCE, Paris, Bois de Vincennes, Pinus, 22. Nov. 1933, leg. R. Kühner (G00058745).

Epitype, selected in Bandini et al. (2022a: 55): GERMANY, Rheinland-Pfalz, Rhein-Pfalz-Kreis, near Dudenhofen, TK25 6616/3, alt. 105 m, sandy soil with Quercus robur, Corylus avellana, Pinus sylvestris, 3 Oct. 2017, leg. D. Bandini (STU SMNS-STU-F-0901730; dupl. priv. herb. D.B. DB3-10-17-6).

Notes

The species was discussed in detail and epitypified in Bandini et al. (2022a). The lectotype was cited as holotype in the latter publication. Kühner (1955) did not cite collection numbers. Other material from the same place and date might exist that would have to be considered as syntype had the lectotypification not been done by Poirier (2016). Epitypes only serve as interpretative types for the material that is cited in the epitypification and not for syntypes. Thus, the typification of I. tarda is settled and less ambiguous than one might conclude from Bandini et al. (2022a).

Typification

Lectotype, designated here (MBT 10015420): FRANCE, near Meaux, Pinus, Oct. 1929, leg. Mauguin. Heim (1931) Le Genre Inocybe, Plate XIX, fig. 6.

Epitype, designated in Bandini et al. (2021c: 1099): GERMANY, Rheinland-Pfalz, Bad Kreuznach, Lützelsoon, near Kellenbach, alt. 400 m, Picea abies, Fagus sylvatica, Pseudotsuga menziesii, 24 Oct. 2015, leg. D. Bandini & B. Oertel (STU SMNS-STU-F-0901532; dupl. priv. herb. D.B. DB24-10-15-3).

Notes

The species was discussed in detail in Bandini et al. (2021c). The here-designated lectotype was listed as holotype there. See under I. geophylla for an explanation of the lectotypification.

Basionym: Agaricus transitorius Britzelm., Ber. Naturhist. Vereins Augsburg 26: 137 (1881).

Typification

Lectotype, designated here (MBT 10015421): GERMANY, Bayern, Krumbach. Britzelmayr (1881), Ber. Naturhist. Vereins Augsburg 26: Abb. 11.

Epitype, selected in Bandini et al. (2022a: 58): NORWAY, Telemark, Kragerø, Kammerfossåsen, Tilia, Fraxinus, Populus forest on clay ground, margin of meadow, under Tilia and Populus, 15 Jul. 2016, leg. T. E. Brandrud TEB-45-16 (O-F-304850).

Notes

The species was discussed in detail in Bandini et al. (2022a). The illustration designated as lectotype here was cited as holotype in the latter publication. A collection by J. STANGL deposited in M (M-0022272) and referred to by Stangl & Kuyper (1985) and Stangl (1989) is labelled as neotype but was apparently never published as type, neither (incorrectly) as neotype nor as epitype. See under I. geophylla for an explanation of the lectotypification.