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Holotype: Western Australia, Walpole, property 1683, Dingo Flat Rd, 34°57’0.000’’S, 116°50’0.000’’ E, in litter under Allocasuarina decussata, Eucalyptus marginata, E. jacksonii and E. guilfoylei, 5 June 1995, N.L. Bougher, K. Syme & M.E. Alosi E5412 (PERTH 07551681). GenBank: OR441039 (ITS).
= R. multicolor Bougher & Syme, Fungi Southern Australia 146 (1998), nom. inval.
Fig. 4. Russula discolor sp. nov. Fresh basidiomata. a. PERTH 07554699. b. PERTH 07551681 (holotype). c. PERTH 07710259. Photo credits: N. Bougher.
Figure 5. Russula discolor sp. nov. (holotype). Hyphal extremities of the pileipellis near the pileus margin (a) and in the centre (b). Scale bar = 10 µm. Drawings B. Buyck.
Figure 6. Russula discolor sp. nov. (holotype). Hyphal extremities of the stipitipellis from upper (a) and lower part of the stipe (b). Scale bar = 10 µm. Drawings B. Buyck.
Figure 7. Russula discolor sp. nov. (holotype). Elements of the hymenium: a. Spores as observed in Melzer's reagent. b. Basidia. c. Basidiola. d. Gloeocystidia on sides of lamellae with contents as observed in Congo red in two cells. Scale bar = 10 µm, but only 5 µm for spores. Drawings B. Buyck.
Diagnosis: Russula discolor is characterised by its occurrence in the eucalypt forests of southwestern Australia, its rapidly discolouring reddish pileus becoming mottled with red, maroon, cream, yellow and grey or greenish grey in variable proportions, by its unchanging white stipe surface and context, the mild taste and the yellow to cream, equal gills producing an egg yellow spore print.
Etymology: in reference to its variegated colours and to the fact that different fruiting bodies may be of different dominant colouration
Pileus 40-85(120) mm broad, convex with incurved margin, soon flat-convex with plane margin, then expanding to flat or with a shallow central depression; margin thin, smooth or very shortly translucent-striate; surface when young mainly red (8D-E8, 9-10C7, 10E5, 10-11D8), soon discolouring and becoming mottled with various colours - e.g. pale pastel yellow (2A2), cream (3-4A4), yellow (2-5A4), pink (10D7, 11C6), orange (4A6), tan brown (7C3), maroon (11F7), violet brown (11E4), grey (2B3-3B2-3) or greyish green (5C3); mature caps may vary from being heavily mottled with a mixture of these colours to remaining mainly red or, on the contrary, ending up completely discoloured and mainly cream, but usually retaining some red traces at the center; sticky when young in damp conditions, readily drying to become smooth but dull, often with a distinct white bloom or scurfy aspect (particularly under hand lens), concentrically wrinkled if excessively drying out (air-dried specimens show this feature). Lamellae adnate or subdecurrent, becoming narrowly adnexed when mature, equal in length or with occasional shorter lamellulae, up to 9 mm high, normally disposed (meaning ca. 1L/mm), fragile, some anastomoses present, cream (2A3-3A2) at first, then deeper yellow-cream (4A4-4A5), unchanging when bruised; lamella edge smooth, entire, becoming more yellow than sides with age. Stipe 35-50(–60) x 10-25 mm, cylindric or sometimes slightly bulbous at the base, white, smooth yet finely and longitudinally rugulose, not bruising in age or when injured; inside solid then stuffed with soft tissue, sometimes developing 1-2 cavities. Veil absent. Context white, unchanging. Basal mycelium white, sparse. Odour not distinctive. Taste mild. Spore deposit butter to egg yellow (4A4-5).
Spores subglobose to broadly ellipsoid, (7.5–)7.9–8.20–8.5(–8.7) x (6.5–)6.7–7.05–7.4(–7.9) µm, Q = (1.06–)1.11–1.16–1.22(–1.29) [E5412 Holotype] (7.1–)7.7–8.21–8.7(–9.2) x (6.2–)6.5–6.88–7.3(–7.7) µm, Q = (1.06–)1.12–1.20–1.28(–1.38) [E5000]; ornamentation up to 1 µm high, subreticulate to reticulate, heavily crested with crests composed of linearly aligned, obtuse to conical warts in a ramifying pattern that is usually transversely oriented to the spore axis, with rare and short linear connections or isolated warts; suprahilar spot moderately amyloid, becoming greyish but not black in Melzer’s reagent, not elevated, often just verruculose or only partly amyloid; apiculus well-developed. Basidia 29–42(–52) x 9–11(–12) µm, mostly short, hardly clavate, often somewhat narrowed in the middle, hyaline, sterigmata up to 6(–9) µm long, 4-spored. Marginal cells not observed; the gill edge being mainly composed of cheilogloeocystidia. Hymenial gloeocystidia abundant on sides of lamellae, 55–85 x 8–11(–15) µm, arising from the lamellar trama, hardly emergent above the basidia, thin–walled, slender, fusiform to cylindrical, with rounded or mucronate apex, with yellowish or greyish granular contents; those near or on the lamallar edges very abundant and slightly smaller, 40–70 x 5–10 µm, with rounded or tapering apex, mainly hyaline, originating in the hymenium, subhymenium or trama. Lamellar trama predominantly composed of sphaerocytes up to 50 µm broad, mixed with few hyphae 2-5 µm broad and oleiferous hyphal fragments. Subhymenium pseudoparenchymatic, composed of tightly packed, irregular cells up to 12 µm broad. Pileipellis hyaline in KOH, pinkish in water, orthochromatic in Cresyl blue, two-layered and composed of a strongly developed pseudoparenchymatic suprapellis covering a subpellis composed of gelatinised hyphae 2–4 µm broad. Suprapellis arranged more or less in a palisade from pileus center to pileus margin, composed of long, multi-celled (up to 10 cells or more outside the pileus center), strongly ramifying, diverticulate, thin-walled hyphal endings that are composed of often strongly and irregularly inflated cells that are globose, barrel-shaped, clavate, ellipsoid or variously shaped, 3–25 µm diam., many with lateral branches or outgrowths; the terminal cell often also very irregular, particularly in the pileus center, but frequently narrowing upward, often (very) short, mostly 12–36 µm long and less than 5 µm wide, some up to 10 µm wide, bottle-shaped. Primordial hyphae, pileogloeocystidia and cystidioid hyphae absent. Stipitipellis less developed; hyphal endings more slender, with terminal cells often short, narrow or irregularly sinuate, toward the stipe base with more irregular, nodulose-sinuate terminal cells as in pileipellis. Caulocystidia not observed. Clamp connections absent in all tissues.
Habitat: on soil amongst litter in Karri (Eucalyptus diversicolor) and Tingle (E. jacksonii & E. guilfoylei) forests, and Jarrah (E. marginata) - Marri (Corymbia calophylla) forests in south-west Australia. Usually in small groups, can be locally abundant.
Additional material examined:
AUSTRALIA. Western Australia, Walpole-Nornalup National Park: Hilltop Road, to Giant Tingle Tree track, in litter under Eucalyptus diversicolor, E. jacksonii and Allocasuarina decussata, 35°1’0.000’’ S, 116°40’0.000’’ E, 12 June 1988, leg. R.N. Hilton E4068 (PERTH 07550308); ibid., on track between Gully Rd and Giant Tingle Tree, 34°59’0.000’’S, 116°47’0.000”E, 05 June 1992, N.L. Bougher & K. Syme E4743 (PERTH 07554648); ibid., Knoll Drive walk, Walpole, in litter under Corymbia calophylla, Eucalyptus diversicolor, E. marginata, Agonis flexuosa and Allocasuarina decussata, 34°59’0.000’’S, 116°43’0.000”E, 06 June 1992, Leg. N.L. Bougher & K. Syme E4746 (PERTH 08792275); ibid., same habitat, 06 June 1992, leg. K. Syme & N.L. Bougher E4748 (PERTH 07554699); ibid., under Eucalyptus diversicolor and E. calophylla, 11 June 1993, N.L. Bougher & K. Syme E5000 (PERTH 07652631). Western Australia, Hull Rd, off North Walpole Rd, near Walpole, under mixed 30 year old rough & smooth barked Eucalyptus species, Agonis parviceps and Acacia pentadenia, 34°53’0.000’’ S, 116°41’50.000’’ E, 04 June 1992, K. Syme & N.L. Bougher E4654 (PERTH 07710259). Western Australia, Mount Shadforth Reserve, Denmark area, 34°58’0.000’’ S, 117°16’0.000’’ E, under Eucalyptus diversicolor, 23 April 1995, leg. K. Syme KS786/95/E5615 (PERTH 07676336, "Morphological details of this collection are described and illustrated in Bougher & Syme 1998 as Russula aff. cyanoxantha"). Western Australia, Amphion Forest Block: Murray River Rd, Dwellingup, 32°47’0.000’’ S, 116°11’0.000’’ E, under unburnt Eucalyptus marginata & Corymbia calophylla, 11 June 1998, leg. N.L. Bougher & S.Q. Bolsenbroek E5943 (PERTH 07605056) & E5949 (PERTH 07605102); ibid., same habitat, 14 June 1998, S.Q. Bolsenbroek E5961 (PERTH 07605226); ibid, under E. marginata, 18 July 1996, N.L. Bougher s.n. (PERTH 07711980). Western Australia, Alcoa Mine, Nettleton Rd, 32°34’0.000’’ S, 116°4’0.000’’ E, under Eucalyptus marginata mixed forest, 14 June 2000, leg. N.L. Bougher s.n. (PERTH 07676336).
Fig. 4. Russula discolor sp. nov. Fresh basidiomata. a. PERTH 07554699. b. PERTH 07551681 (holotype). c. PERTH 07710259. Photo credits: N. Bougher.
Figure 5. Russula discolor sp. nov. (holotype). Hyphal extremities of the pileipellis near the pileus margin (a) and in the centre (b). Scale bar = 10 µm. Drawings B. Buyck.
Figure 6. Russula discolor sp. nov. (holotype). Hyphal extremities of the stipitipellis from upper (a) and lower part of the stipe (b). Scale bar = 10 µm. Drawings B. Buyck.
Figure 7. Russula discolor sp. nov. (holotype). Elements of the hymenium: a. Spores as observed in Melzer's reagent. b. Basidia. c. Basidiola. d. Gloeocystidia on sides of lamellae with contents as observed in Congo red in two cells. Scale bar = 10 µm, but only 5 µm for spores. Drawings B. Buyck.
Notes:
The description of the new R. discolor depicts a species with a variable pileus colour. In general terms, it could be described as an overall reddish species that discolours easily and rapidly, while developing different tinges. Also the colour of the lamellae changes distinctly between young and mature fruiting bodies. Notwithstanding this important colour variation, the species is considered in Australia to be ‘easily recognizable in the field’ (Richardson 2011, Bougher & Syme 1998). This was confirmed by our sequencing results because all of the received collections (see studied material) produced identical ITS sequences. It seems to be widespread in southwestern Australia where it is exclusively associated with various Eucalyptus species that compose the Karri (Eucalyptus diversicolor), Tingle (E. jacksonii & E. guilfoylei) and Jarrah (E. marginata) - Marri (Corymbia calophylla) forest stands. We can also confirm here that the voucher that served for the illustration and description in Bougher & Syme (1998) of R. aff. cyanoxantha / R. multicolor [as listed on p. 380 of that work] was successfully sequenced and represents indeed R. discolor.
Russula wielangtae, described from Tasmania and equally a eucalypt associate, is definitely close (ITS similarity between 97-98%) to the here newly described R. discolor and shares with it a very similar spore ornamentation, a similar composition of the pileipellis and even a similar overall colour, although less variable. The presence of ‘pileocystidia’ was considered ‘doubtful’ in the original description of R. wielangtae and the legend of fig. 74 in Adamčík et al. (2019), mentioning twice ‘pileocystidia’, probably needs correction as none of the hyphal terminations in the pileipellis of these Oceanian species has the typical contents of the generally accepted definition of gloeocystidia in the genus. Russula wielangtae might also occur on the Australian continent as suggested by the collection MEL 2238232 (see Fig. 3a,b). The ITS sequence for this collection was deposited in GenBank associated with the name R. wollumbina Grgur. and still represents the single deposited ITS sequence under that name (Lebel & Tonkin 2007). Unfortunately this sequence seems of unsufficient quality, lacking more than 70 base pairs at the start of ITS1 and having several ambiguous base pairs further down the sequence. The specimen was identified and collected by J. E. Tonkin in eucalypt woodland in southeastern Australia near Mt Richmond (https://biocache.ala.org.au/occurrences/9063ba82-91d0-4c74-9e1f-329ed608b9db). Tonkin's identification was questioned by Adamčík et al. (2019) who assumed that it rather represents their newly described R. wielangtae (we agree on the basis of the ITS sequence similarity) as the spores of the type specimen of R. wollumbina were described and illustrated as having much more isolated warts and scarcely any ridges (Grgurinovic 1997), a feature that was confirmed for the type in Adamčík et al. (2019).
There are many more collections reported as R. wollumbina in the ‘Atlas of Living Australia’ (https://www.ala.org.au), but none of these have been sequenced. The fact that the type of R. wollumbina is based on a collection by Cleland (1934) originally identified as R. xerampelina, a species often reported from Australia in the past (e.g.: Dickinson & Lucas 1979, Gill & Ashton 1971, Shepherd & Totterdell 1988, Sinnott 1976) seems to be responsible for a misleading shortcut. Indeed, certain websites on Australian mushrooms automatically seem to assume that all mentions of R. xerampelina in Australia correspond now to R. wollumbina, which is absolutely not the case. An interesting and recent find that might correspond to R. wollumbina is a description by P. Leonard for a specimen identified as such and uploaded on https://qldfungi.org.au/wp-content/uploads/2022/02/Russula-wollumbina.pdf. This specimen indeed shows some overall resemblance to close relatives of R. xerampelina, the earlier interpretation of R. wollumbina. Leonard’s description mentions pale gills that produce a whitish or very pale cream spore print (IIa following the Romagnesi 1967 scale), not the ochre or yellow spore print observed for R. wielangtae or R. discolor, respectively, and stresses the often strongly tapering stipe, two characters that match the original interpretation of Cleland (1934, as ‘R. xerampelina’). However, the spores of Leonard’s collection are described as having strongly amyloid ridges, which seems in contradiction with the spore description given by Grgurinovic (1997) for the type specimen of R. wollumbina. Leonard’s interpretation of R. wollumbina suggests a very different species compared to the here newly described R. discolor which has more variegated colours, much darker spore print and a cylindrical to slightly bulbous stipe. The stipe of R. discolor also lacks the pinkish tints observed on nearly all of the other species that are discussed here. A correct interpretation of R. wollumbina remains therefore problematic for the moment.
The only other species that has been placed in the /wielangtae lineage in recently published phylogenies was identified as R. atroviridis Buyck from New Zealand (Adamčík et al. 2019, Cooper 2021). It differs already from both other Oceanian species in this lineage because it associates not with eucalypts, but principally with Nothofagus. Our phylogeny (Fig. 3b) suggests that there are possibly two different, but very close phylogenetic entities. Although considerable effort has been made to obtain sequences for most of the Russula species of New Zealand (Cooper & Leonard 2014, Cooper 2021), there remains an urgent need for better and more detailed descriptions, particularly for the type specimens. We, therefore, provide below our own microscopic examination of the isotype of R. atroviridis. As discussed below, the results of this re-examination raised more questions.