Two new species in the Lulesia fallax complex (Entolomataceae, Agaricales) from Europe (Fennoscandia and Switzerland), Lulesia neofallax comb. nov., new records of recently described species of Lulesia, and notes on Clitocybe alachuana

Two new species in the Lulesia fallax complex (Entolomataceae, Agaricales) from Europe (Fennoscandia and Switzerland), Lulesia neofallax comb. nov., new records of recently described species of Lulesia, and notes on Clitocybe alachuana | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Two new species in the Lulesia fallax complex (Entolomataceae, Agaricales) from Europe (Fennoscandia and Switzerland), Lulesia neofallax comb. nov., new records of recently described species of Lulesia, and notes on Clitocybe alachuana Alfredo Vizzini, Giovanni Consiglio, Tapio Kekki, Mauro Marchetti, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7232140/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 21 Nov, 2025 Read the published version in Mycological Progress → Version 1 posted 6 You are reading this latest preprint version Abstract Lulesia fallacioides (from Finland) and L . parvifallax (from Switzerland) are described as new species, based on both a morphological and molecular approach, in the L . fallax complex ( Lulesia subgen. Paraclitopilus ), a morphology-delimited aggregate of species sharing white basidiomes and a bitter context. Compared to L . fallax , the former differs mainly by wider spores and a gelatinized pileipellis and the latter by smaller basidiomes and shorter spores. The recently described Clitocella neofallax from China is here combined in Lulesia . New collections of L . colorata and L . solaris made in Italy and Finland have allowed us to better understand the distinctive features and/or to extend the distribution area of ​​these recently established species. The presence of L . fallax , L . mundula and L . obscura in Finland is here molecularly confirmed for the first time. Finally, a full morphological description of L. alachuana , a North American species so far described from Florida, is provided based on ancient (holotype included) and recent collections. Basidiomycota Agaricomycetes Tricholomatineae Clitocella Rhodocybe cryptic species multigene analysis taxonomy Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction The genus Lulesia was established by Singer ( 1970 ) for accommodating two species previously considered as part of Armillariella Singer (Singer 1951 ; Singer and Digilio 1951 ), viz. A . densifolia Singer (type of Lulesia , from Tucumán, Argentina) and Clitocybe alachuana Murrill (from Florida, USA). Singer ( 1970 ) differentiated Lulesia from Armillariella by having a zonate pileus, very narrow lamellae, a bitter taste, smaller spores, a trichodermic pileus covering, and a terricolous/humicolous habitat. Lulesia densifolia spores were described as smooth in some mounting media (methylene blue, carmine-acetic acid, and Melzer’s reagent) but appearing slightly rounded-angular and nodulose in aqueous NH 3 and KOH in light microscopy (Singer and Digilio 1951 ; Singer 1970 , 1986 ). According to Singer and Digilio ( 1951 ) and Singer ( 1986 ) this slight irregularity of the spore surface would be reminiscent of the spores of the species in genus Rhodocybe Maire. Baroni ( 1981 ) and Bigelow ( 1982 , 1985 ), after studying the holotype collection of Clitocybe alachuana , considered it a later synonym of Rhodocybe mundula (Lasch) Singer, and this conclusion was also followed by Singer ( 1986 ). The species recognized in Lulesia were recently supplemented by Lechner et al. ( 2006 ) with another new species from Argentina, L . lignicola B.E. Lechner & J.E. Wright. No further collections of L. densifolia apart from the original ones were reported until recent findings in the Dominican Republic (Angelini and Contu 2012 ). Based only on morphological data, Lulesia was placed by Singer ( 1970 , 1986 ) in subtribe Omphalineae Singer (tribe Clitocybeae Fayod, family Tricholomataceae R. Heim ex Pouzar), by Lechner et al. ( 2006 ) and Agerer ( 2018 ) in Tricholomataceae s.l., and by He et al. ( 2019 , 2024 ) in incertae sedis Agaricales Underw. and Entolomataceae Kotl. & Pouzar, respectively. Recently, based on multigene analyses, Vizzini et al. ( 2024 ) have highlighted that L. densifolia clustered within the genus Clitocella Kluting, T.J. Baroni & Bergemann, a genus (typified by Agaricus popinalis Fr.) segregated from Rhodocybe s.l. ( Entolomataceae ) (Kluting et al. 2014 ) and recently monographed at European level by Vizzini et al. ( 2023 ). This indicated Lulesia as a priority synonym of Clitocella . Three distinct subgenera are currently recognized in Lulesia relied on type of hymenophoral trama, basidiome colours and reactions of basidiome surface to KOH: subgen. Lulesia , Paraclitopilus (Vizzini & Consiglio) Vizzini & Consiglio, and Rhodopleurella (Vizzini & Consiglio) Vizzini & Consiglio (Vizzini et al. 2024 ). Lulesia species typically produce a gymnocarpic centrally stipitate to pleuropodal ( L . termitophila (T.J. Baroni & Angelini) T.J. Baroni & Angelini), clitocyboid basidiome, which exhibits a small to large (≤ 120 mm) pileus that can range from white to greyish, grey-brown or purplish grey in colour, usually covered with a white bloom, long-decurrent narrow lamellae with pinkish tinges at maturity, a cylindrical glabrous to pubescent, floccose or fibrillose stipe usually with white rhizomorphs at the base, context usually with farinaceous to cucumber-like smell and taste bitter, surfaces of the dried basidiomes reddening or not in KOH, spore-print incarnate-pink, hymenophoral trama regular (consisting of parallel cylindrical hyphae) or irregular (composed by interwoven hyphae), hymenial cystidia usually lacking, and cyanophilic, inamyloid, non-dextrinoid, thin-walled basidiospores with obscure or distinct undulating pustules or minute bumps (Kluting et al. 2014 ; Vizzini et al. 2023 , 2024 ). To date, a total of 17 species of Lulesia and two uncombined Clitocella species have been validly published based on the records from the Index Fungorum database ( www.indexfungorum.org , accessed 21 January 2025). Lulesia has been reported in Asia, Europe, North, South and Central America, and can be observed in various ecosystems, including forested and grassland systems, as well as on dunes or in association with hind dune trees, from temperate to subtropical/tropical areas (Angelini and Contu 2012 ; Kluting et al. 2014 ; Baroni et al. 2020 ; Jian et al. 2020a ; Mao et al. 2022 ; Vizzini et al. 2023 , 2024 ; Liu et al. 2025 ; Xiao et al. 2024 ). Its species are presumably saprotrophic (Vizzini et al. 2023 , 2024 ), usually terrestrial but C . lignicola is reported on dead wood of dicotyledonous trees (Lechner et al. 2006 )d termitophila grows on arboreal nests of neotropical termites (Baroni et al. 2020 ). The aim of the present work was to describe two new white Lulesia species from Europe morphologically closely related to L . fallax , by using both a traditional (morphological) and multilocus molecular approach. Additionally, a full morphological description of the north American species Clitocybe alachuana (= L . alachuana (Murrill) Singer) based on ancient and modern collections, and notes on the recently established species L . colorata and L . solaris (Mao et al. 2022 ; Vizzini et al. 2023 ), are provided. Materials & Methods Morphological studies Macroscopic morphological features were studied in fresh specimens. Codes for colours follow the Methuen Handbook of Colour (Kornerup and Wanscher 1984). The following abbreviations are employed: L = number of lamellae reaching the stipe; l = number of lamellulae between each pair of lamellae. Microscopic structures were examined in dried material using different mounting media: water, L4 (Clémençon 1972), Melzer’s reagent, ammoniacal Congo red, phloxine, and Cotton blue. Dried pieces of the samples were rehydrated in water and mounted in L4. All microscopic measurements were carried out with a Nikon Eclipse 80i microscope, using immersion oil at ×1000 magnification. Basidiospore measurements were made by capturing images of a single visual field with multiple spores (taken from lamellar squashes of exsiccated material of mature specimens) which were then measured using the DS-L1 Nikon camera control unit. Basidiospore dimensions excluded the hilar appendix and are given as (a)b– c –d(e), where (a) = minimum value, b = (average minus standard deviation), c = average , d = (average plus standard deviation) and (e) = maximum value; Q = (minimum−) average minus standard deviation – average – average plus standard deviation (−maximum) of ratio length/width; V = (minimum−) average minus standard deviation – average – average plus standard deviation (−maximum) of the volume. The approximate spore volume was calculated as that of an ellipsoid (Gross 1972; Meerts 1999). The notation [n/m/p] indicates that measurements were made on ‘n’ randomly selected spores from ‘m’ basidiomes of ‘p’ collections. A minimum of 30 spores were measured for each collection. Q indicates the quotient of length and width of the spores in side view. The width of the basidia was measured at the widest part, and the length was measured from the apex (sterigmata excluded) to the basal septum. Photographs of microscopic features were taken using a Nikon DS 5M digital camera with a preset resolution of 2560 × 1980 pixels connected to a Nikon Eclipse 80i microscope with both brightfield and interferential contrast optics and saved in TIFF format. They were then cleaned using Adobe Photoshop CC 2019 version 20.0.10, in some cases replacing, through the Magic Wand Tool and with the appropriate tolerances, the background colour of the photograph with an alternative uniform background to highlight the morphology of the microscopic characters, which maintained their original size and hue. In other cases, the best individual spores from microscopy photos were copied using the Polygonal Lasso Tool or alternatively the Magnetic Lasso Tool of Adobe Photoshop CC 2019 version 20.0.10, onto a new monochrome sheet, with the same resolution (2560 × 1980 pixels) to maintain the spore size and original colours as previously acquired. Macro- and microchemical testing of pigments were performed using basic solutions (5% KOH and 10% ammonia, separately). Chemical spot tests were performed on pileus surface, lamellae and stipe of fresh and/or dried basidiomes using 3% and 10% KOH, following Baroni (1978, 1981). Fungarium acronyms follow Thiers (2025) except that ANGE and T. Kekki refer to the personal herbarium of Claudio Angelini and Tapio Kekki, respectively. DNA extraction, amplification, and sequencing Total DNA was extracted from seventeen dry specimens (Fig. 1, Tab. 1) employing a modified protocol based on Murray and Thompson (1980). PCR (Mullis and Faloona 1987) reaction conditions (cycles and annealing temperature) follow Alvarado (2010, 2012). The primers ITS1F and ITS4 (White et al. 1990; Gardes and Bruns 1993) were used to amplify the ITS rDNA nuclear region, LR0R and LR5 (Vilgalys and Hester 1990; Cubeta et al. 1991) for the LSU rDNA nuclear region, EF1-983F, EF1-1567R and EF1- 2218R (Rehner and Buckley 2005) for the translation elongation factor 1-alpha ( TEF1-1α ) gene, and bRPB2-6F2 (reverse of bRPB2-6R2), and bRPB2-7R2 for the RNA polymerase II second largest subunit ( RPB2 ) gene (Matheny et al. 2007). PCR products were checked in 1% agarose gels, and amplicons were sequenced with one or both PCR primers. Sequences were corrected to remove reading errors in chromatograms. Phylogenetic analyses Two different dataset/alignments were built. 1) A Lulesia -dataset aimed to provide an accurate view of the position of the newly sequenced collections within this genus was prepared. It included sequences of five different loci (nrITS, nrLSU, RPB2 -exons, TEF-1α -exons and ATP6 ) from all the species recognized in Vizzini et al. (2023, 2024) supplemented with those present in Xiao et al. (2024) and Liu et al. (2025), and the most closely related sequences selected from public databases as the International Nucleotide Sequence Database Collaboration public database (INSDC/GenBank https://www.ncbi.nlm.nih. gov/genbank/, Arita et al. 2021), UNITE (https://unite.ut.ee/), and BOLDSYSTEMS (http://www.boldsystems.org/), via BLASTn algorithm (Altschul et al. 1990). Clitopilus prunulus was employed as outgroup taxon following Baroni et al. (2020), Mao et al. (2022) and Vizzini et al. (2023, 2024). 2) A nrITS dataset focused on Lulesia subgen. Paraclitopilus and including also all environmental sequences was prepared, with C . popinalis as outgroup taxon. All the sequences employed are listed in Table 1. The combined and ITS alignments used for the phylogenetic analyses are deposited in Figshare (https://doi.org/10.6084/m9.figshare.29654816). Sequences first were aligned in MEGA 6.0 (Tamura et al. 2013) with its MUSCLE (Edgar 2004) applications and then realigned manually as needed to establish positional homology. The matrices were analysed by using the Bayesian Inference (BI) and the Maximum Likelihood (ML) criteria through the MESQUITE 3.81 (Maddison and Maddison 2023) software by which were obtained the .nex and .phy files. The .nex files were loaded into MrBayes v. 3.2.7a (Ronquist et al. 2012) of the CIPRES Science Gateway v. 3.3 platform (Miller et al. 2010) and Bayesian analyses were performed with the nrITS–nrLSU– RPB2 – TEF-1 α - ATP6 partitioned alignment and the ITS alignment (GTR+G+I model, two simultaneous runs, six chains, temperature set at 0.2, sampling every 1 000 generations) until reaching convergence parameters (standard deviation less than 0.01 and PSRF (Potential Scale Reduction Factor) (Gelman and Rubin 1992) equal to 1), after 3.17 M generations and 13.45 M generations, respectively. As required from the procedure, 25 % of the trees, those of the initial stretch and those of the final tail, were ‘burned’. A full search for the best-scoring maximum likelihood tree was performed loading the .phy files of the multilocus alignment and ITS alignment into the RAxML v. 8.2.12 program (Stamatakis 2014) using the standard search algorithm with same partitions and 1 000 bootstrap cycles according to the GTR + GAMMA model. The trees in .tre format were read with the software SEAVIEW v. 4 (Gouy et al. 2010) and saved in a vector format for printing. The significance threshold was set ≥ 70% for bootstrap proportions (BP) and ≥ 0.95 for posterior probabilities (PP). Results Molecular data The topology of the best tree from the ML analysis and 50 % majority-rule consensus tree from the BI analysis are essentially identical, so only the ML tree is presented here, with support values from both analyses (Bootstrap (BS) ≥ 70 % and/or Posterior Probabilities (PP) ≥ 0.95) (Figs 1–2). The final combined multigene nrITS–nrLSU– RPB2 – TEF-1α – ATP6 Lulesia data matrix encompassed a total of 339 sequences (including 27 newly generated, 303 from INSDC/GenBank and 9 from UNITE) (100 ITS, 66 LSU, 84 RPB2 , 51 TEF-1α , 38 ATP6 ) from 132 samples of 21 taxa. The alignment is 3387 bp long (gaps included). The nrITS Lulesia subgen. Paraclitopilus data matrix encompassed a total of 34 sequences (including 7 newly generated, 22 from INSDC/GenBank and 5 from UNITE) from 34 samples of 8 taxa. The alignment is 604 bp long (gaps included). Except for L . termitophila , all the other species of Lulesia included in the present multigene analysis clustered together forming a monophyletic lineage (BP = 100%, PP = 1.00). Lulesia termitophila was sister to all the other species of Lulesia (BP = 100%, PP = 1.00) (Fig. 1). Three major clades are recognized in Lulesia , corresponding to Lulesia subgen. Lulesia (BP = 92%, PP = 1.0), subgen. Paraclitopilus (BP = 92%, PP = 1.00) and subgen. Rhodopleurella , as previously highlighted by Vizzini et al. (2023, 2024). Lulesia encompasses nineteen monophyletic species-rank clades corresponding to fifteen already described species, two new species and two Lulesia sp. ( Lulesia sp. 1 and Lulesia sp. 2) (Fig. 1). All these species clades with more than one sequence were strongly supported. The two new species (described here as L . fallacioides and L . parvifallax ) and Lulesia sp. 2 belong to Lulesia subgen. Paraclitopilus which also includes L . blancii , L . fallax , and L . neofallax (Fig. 1). Lulesia fallacioides , L . parvifallax and Lulesia sp. 2 (as Lulesia sp.) are clearly supported as distinct taxa also in the nrITS analysis focused on Lulesia subgen. Paraclitopilus (Fig. 2). Lulesia sp. 2 consists of two environmental sequences from USA (Utah) and two basidiomatal sequences (China-Xizang and USA-California). Morphological data will be needed to formally describe the clade as a new species. Taxonomy Lulesia subgen. Paraclitopilus (Vizzini & Consiglio) Vizzini & Consiglio, in Vizzini, Alvarado, Consiglio, Angelini & Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 8 (2024) [2023] Basionym: Clitocella subgen. Paraclitopilus Vizzini & Consiglio, Persoonia 50: 146 (2023) Type: Omphalia fallax Quél., C. r. Assoc. Franç. Avancem. Sci. 24(2): 617 (1896) [1895] Habit like that of the centrally-stipitate Clitopilus species, pileus white to cream coloured under a white pruinose covering, surfaces of both fresh and dried basidiomes not staining grey or black when bruised or in age and with a negative KOH reaction, hymenophoral trama regular, basidiospores broadly ellipsoid, ellipsoid to oblong, amygdaliform, obscurely angular, often adhering (grouping) in tetrads, and sporal Q on average usually exceeding (1.20–)1.30. Lulesia fallacioides Kekki, Vizzini, Consiglio, M. Marchetti & Kytöv., sp. nov. Figs. 3a–c, 5 MycoBank MBXXXXXXX Etymology: fallacioìdes = from the root fallac- of fallax (fallac-is) (Latin) and the suffix -ìdes relating to Greek ειδος (éidos), which means “external aspect, form, semblance”, therefore indicating resemblance to Lulesia fallax Diagnosis : It differs from L . fallax by an ixocutis type of pileipellis, wider basidiospores (4.7 μm on average), and association with Picea abies forests Holotype (here designated): OULU:GAJ.17962 Pileus 10–50(–80) mm diam., convex, then applanate and usually depressed in the centre, often irregular when old. Margin involute when young, then straight, usually strongly wavy and undulating; surface dry, smooth to slightly tomentose, not hygrophanous, not translucently striate, white, sometimes with slight cream or ochre tinges in the centre when old. Lamellae L = 20–45, l = (2–) 3–5, crowded, arcuate-decurrent, 2–4 mm broad, sordid white, finally tinged with pink, pinkish puff to clay-puff tinges when dry, with a smooth concolorous edge. Stipe 8–26 × 2–7 mm, cylindrical, often tapering downwards, subclavate at the base, solid, white, usually with white rhizomorphs at the base. Contex solid, white. Smell not distinctive, taste bitter. No part of the basidiome staining black. Spore-print pale cream pink. Chemical spot-test reactions KOH on dried basidiomes surfaces (pileus, lamellae and stipe) produces no reaction (negative). Basidiospores (4.81–)5.9– 6.5 –7.2(–8.45) × (3.59–)4.4– 4.7 –5.1(–5.70) µm [175/3/3], Q = (1.10–)1.27– 1.38 –1.50(–1.79), V = (37.9–)59.2– 76.8 –94.5(–138) μm 3 , subamygdaliform in side view, ovoid-ellipsoid in frontal view, minutely angular in polar view (7–10 facets), weakly nodulose-pustulate to verrucose in all views, rounded to slightly attenuated apically, often adhering in tetrads, thin-walled (no sclerospores), hilar appendage evident, up to 1 μm long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cyanophilic, pale yellow in water, yellow in Melzer’s. Basidia 18–30 × 7–10 μm, cylindrical to clavate-cylindrical, usually with guttulate contents, mainly tetrasporic, rarely bisporic, sterigmata up to 4 μm long. Subhymenium 20–30 μm thick, consisting of textura intricata short elements, 2.5–5 μm wide. Hymenophoral trama regular to subregular of parallel to slightly intervowen cylindrical, thin-walled hyphae, hyaline, 3–10 μm wide, sometimes inflated at septa. Hymenial cystidia absent. Pileipellis an ixocutis of thin-walled, 3–8 μm wide, subparallel to loosely intertwined cylindrical hyphae, sometimes inflated at septa, faintly yellowish; terminal ascending (erect) elements rare to frequent, immersed in a gelled matrix, mostly in clusters, sinuous, subcylindrical with rounded apex, 2–4 μm wide, often multi-articulated into multiple segments; pigment absent to pale yellow, parietal or rarely minutely incrusting some hyphae. Subpellis of subparallel to loosely intertwined 3–10 μm wide hyaline hyphae. Thromboplerous hyphae not observed. Stipitipellis consisting of parallel, faintly yellowish, 2–4 μm wide cylindrical repent hyphae, occasionally producing clusters of erect, clavate, subglobose to cylindrical (and then multi-articulate) 4–8 µm wide hyaline caulocystidioid hyphae (elements). Stipititrama of hyaline, short cylindrical, slightly thick-walled (wall up to 0.5 μm thick), 3–7 μm wide hyphae. Clamp connections absent. Habitat and distribution : Boreal forests of Picea abies , gregarious on needle litter, August-October (European collections), so far known from Finland, Sweden and North America (USA, Oregon). Material examined : Finland , Kainuu, Paltamo, Kontiomäki, mesic Picea abies forest, 14 September 2008, leg. I. Kytövuori , H6032677 (H); Suomussalmi, Lohivaara, herb-rich Picea abies forest on needle litter, 29 August 2019, leg. T. Kekki (3988) & T. Helo , OULU:GAJ.11332; Koillismaa, Taivalkoski, Katajavaara, old mesic Picea abies forest with damp grass-herb depressions, on needle litter, 02 September 2008, leg. I. Kytövuori , H6082249 (H); Perä-Pohjanmaa, Kemijärvi, Pyhätunturi, fairly old Picea abies forest, 27 August 2008, leg. I. Kytövuori , H6032673 (H); Rovaniemi, Kylmäoja, mesic Picea abies forest by a stream, on needle litter, 04 October 2023, leg. T. Kekki (6650) , OULU:GAJ.17963; Tervola, Raemäki, calciferous Picea abies forest, on needle litter, 04 October 2023, leg. T. Kekki (6670) , OULU:GAJ.17962 ( Holotype , Isotype AMB 20536); ibid. 04 October 2023, leg. T. Kekki (6666) , OULU:GAJ.17965; ibid. 04 October 2023, leg. T. Kekki (6668), OULU:GAJ.17964. Sweden , Jämtland, Åre, mesic Picea abies forest, 23 August 2006, leg. M. Toivonen & I. Kytövuori , H7001634 (H). Notes – The species grows gregarious, in autumn, in the thick needle litter under Picea abies . It is probably circumpolar as there are collections from Finland, Sweden, and North America. It is rare and prefers old growth Picea forests. Lulesia fallacioides can be confused with Leucopaxillus alboalutaceus (F.H. Møller & Jul. Schäff.) F.H. Møller, Ripartites P. Karst. spp. and centrally-stipitate Clitopilus (Fr. ex Rabenh.) P. Kumm. species, but its basidiospore structure is diagnostic. Lulesia fallax is morphologically very similar, but it grows usually in deciduous forests and grasslands, has a xerocutis (versus an ixocutis), and narrower oblong basidiospores (4.1 μm and Q = 1.60 versus 4.7 μm and Q = 1.38, on average) (Vizzini et al. 2023). The Swedish collection named Rhodocybe fallax in von Bonsdorff et al. (2014), due to its habitat, viscid pileus and basidiospore shape is surely attributable to L. fallacioides . The short description of R . fallax included in Funga Nordica by Noordeloos (2012), with basidiospores 5–7 × 3–4 µm, pileus 10–40 mm, stipe 2–7 mm wide, in deciduous woods, scrubs, calcareous sand dunes, in litter under Syringa , Alnus , indicates that the true L . fallax is present in Scandinavia. Our analyses (Figs. 1–2) show that based on molecularly confirmed collections, L . fallax is present in Nordic areas as Finland (H6032675, Fig. 4a) and Estonia. The Chinese L . neofallax (see below) and the Swiss L . parvifallax (see below) are mainly distinguished from L . fallacioides by smaller basidiomes, a xerocutis as pileipellis, smaller basidiospores, and different habitat (Tab. 2, Liu et al. 2024). Lulesia blancii (Maire) Vizzini, Consiglio, P. Alvarado, Angelini & M. Marchetti is a xerophilic species from Mediterranean basin distinguished by a dry alutaceous ochre, chamois, grey-brown pileus, lamellae with evident yellow-ochre tinges, and shorter basidiospores (5.5 µm long on average) (Contu 1999; Eyssartier and Roux 2011; Ivaldi et al. 2023; Vizzini et al. 2023). Lulesia parvifallax Rickmann, Vizzini, Consiglio & A. Gross sp. nov. Figs. 3d–e, 6 MycoBank MBXXXXXXX Etymology : from the Latin adjectives parvus (small) and fallax (false, fallacious), due to its similarity to Lulesia fallax , from which it differs especially by having smaller basidiomes and smaller basidiospores. Diagnosis : It differs from L . fallax by smaller basidiomes and smaller basidiospores. Holotype (here designated): G00586348 Pileus 15–35(–45) mm diam., convex, soon applanate or depressed at center, getting irregular when old. Margin involute when young, later more or less undulating, often crenulate, not striate. Surface smooth or slightly pruinose, dry, white, sometimes with cream tinges, not hygrophanous, often concentrically cracking and showing the darker context. Lamellae L = 25–40, l = 1–3, moderately crowded, decurrent, anastomosing, 1.5–3 mm broad, whitish, later tinged pinkish cream, edge smooth, concolorous. Stipe 10–35 × 2–5 mm, central or slightly eccentric attached, cylindrical, usually tapering downwards, smooth, solid, ivory white to pale ochre beige, often with a tinge of pink towards the base, with white rhizomorphs at the base. Context whitish. Smell not distinctive, taste strongly bitter. No part of the basidiome staining black. KOH on dried basidiome surfaces negative. Spore-print cream pink. Basidiospores (4.7–)5.0– 5.5 –5.9(–6.2) × (3.2–)3.5– 3.9 –4.3(–4.6) µm [60/2/1], Q = (1.26–)1.34– 1.43 –1.55(–1.63), V= (25.7–)34.3– 43.9 –57.9(–66.5), subamygdaliform in side view, ovoid-ellipsoid in frontal view, angular in polar view (5–7 facets), weakly nodulose-pustulate to verrucose in all views, rounded to slightly attenuated or subconical apically, often adhering in tetrads, usually thin-walled, rarely somewhat thick-walled, hilar appendage evident, up to 1 µm long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cyanophilic, pale yellow in water, yellow in Melzer’s. Basidia 22–30 × 6–8 µm, cylindrical to clavate-cylindrical, usually with guttulate contents, mainly tetrasporic, rarely bisporic, sterigmata up to 6 µm long. Subhymenium 30–50 µm thick, consisting of irregular elements, up to 12 µm wide. Hymenophoral trama regular to subregular of parallel to slightly interwoven, cylindrical or inflated, thin-walled, hyaline, 4–14 µm wide hyphae. Hymenial cystidia absent. Pileipellis a xerocutis of thin-walled, 3–6 µm wide, subparallel to loosely intertwined cylindrical hyphae, sometimes inflated at septa, terminal elements inconspicuous, cylindrical, subclavate or subfusiform, with rounded apex, pigment absent or parietal, not incrusting, pale yellow. Subpellis poorly differentiated, of subparallel to loosely intertwined 3–6 µm wide hyaline hyphae. Thromboplerous hyphae not observed. Stipitipellis consisting of parallel, faintly yellowish, 3–5 µm wide cylindrical repent hyphae, terminal elements hardly differentiated. Stipititrama of hyaline, cylindrical, thin-walled, 5–9 µm wide hyphae. Clamp connections absent. Habitat and distribution : Dry grassland on sandy, calcareous soil, accompanied by Pinus sylvestris and Quercus pubescens . So far only known from the type locality. Material examined : Switzerland , Canton of Valais, municipality of Leuk, 572 m a.s.l., sunny, dry area on sandy, calcareous soil; the ground covered with mosses and lichens; nearby vegetation includes Pinus sylvestris, Quercus pubescens , and numerous bushes of Berberis vulgaris ; other fungal species observed in the vicinity were Calocybe carnea (Bull.) Donk , Clitocybe barbularum (Romagn.) P.D. Orton , Entoloma coracis Brandrud, Dima, Noordel., G.M. Jansen & Vila , Infundibulicybe glareosa (Röllin & Monthoux) Harmaja , Lepiota pseudolilacea Huijsman , Lichenomphalia ericetorum (Pers.) Voitk, Thorn & I. Saar and Tulostoma melanocyclum Bres.; 19 October 2024, leg. R. Rickmann (private herbarium no. RR24.326), G00586348 ( Holotype ). Notes – Lulesia parvifallax is so far only known from the type locality in Switzerland. Its precise habitat requirements still need to be clarified through additional collections. In the field, reliable distinction from closely related species is hardly possible. Apart from the habitat, only the rather small basidiomes with slender stipes may serve as an indication of L . parvifallax . However, microscopic differentiation is possible; see also Table 2. Lulesia fallax and L . fallacioides differ by having slightly larger basidiospores, usually longer than 6 µm (Vizzini et al. 2023 and Tab. 2). Furthermore, L . fallacioides has an ixocutis. The recently described L . neofallax from China can be distinguished by very minute stipes (only 1–2 mm wide) and a lower spore quotient (Q = 1.2). So far, this species has not been recorded in Europe (Liu et al. 2025). Despite the morphological similarity of L . parvifallax with the other white-capped species, it is molecularly quite distant from these in the multigene analysis (Fig. 1) while its closest relative is L . blancii (with an ITS sequence similarity of approximately 91%). Lulesia blancii shares with L . parvifallax similar basidiospore dimensions but it has usually significantly larger basidiomes (pileus up to 70(–100) mm broad), darker pileus colours, and is so far only known from Mediterranean regions (Ivaldi et al. 2023; Vizzini et al. 2023). Lulesia neofallax (W.H. Lu, Karun. & S. Tibpromma)Vizzini, Consiglio, comb. nov. MycoBank MBXXXXXXX Basionym: Clitocella neofallax W.H. Lu, Karun. & S. Tibpromma, in Liu et al., Mycology 16(1): 26. (2025) [2024] [MB#571612] In our analysis Clitocella neofallax is phylogenetically part of Lulesia subgen. Paraclitopilus (Figs. 1–2) and it is combined above in Lulesia accordingly. The species is distinguished among the others within the L . fallax complex by gracile basidiomes, pileus 10–30 mm wide, dry, low convex, sometimes infundibuliform, with a shallow depression at the centre, stipe 10‒25 × 1‒2 mm, lamellae with evident yellowish hues even when young, basidiospores (4.0–)4.8–6.3 × (3.5–)4–5 μm, Q= 1.2 and very short basidia, 15‒23.5 × 6‒9 μm (Liu et al. 2025). It is so far known only from China (Yunnan Province, Qujing City, Qujing Normal University) on soil associated with bamboo roots. Lulesia subgen. Lulesia Singer, Fl. Neotrop., Monogr. 3: 16 (1970) Autonym Type: Armillariella densifolia Singer, in Singer & Digilio, Lilloa 25: 72 (1952) [1951] Pileus usually pale grey, grey, dark grey, brown, violaceous black, basidiome surfaces unchanging or turning grey or black when bruised or with age, usually with a positive, reddish, KOH reaction, hymenophoral trama usually irregular (intertwined hyphae), and basidiospores globose, subglobose, broadly ellipsoid to ellipsoid, which are obscurely to clearly angular, and weakly to clearly pustulate. Lulesia alachuana (Murrill) Singer, Fl. Neotrop., Monogr. 3: 17 (1970) Figs. 3i–j, 7–8 Basionym: Clitocybe alachuana Murrill, Proc. Fla Acad. Sci. 7(2/3): 107 (1945) [1944] Holotype : FLAS F-17903, USA, Florida, Alachua, Prairie Creek Hammock, on dead leaves, 15 July 1938, leg. West, Arnold, and Murrill , det. Murrill . Epitype : FLAS F-61088 (Designated by Vizzini, Alvarado, Consiglio, Angelini & Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 5. 2024). Registration Identifier 10017392. MycoBank MBT10017392 Pileus 20–80 mm diam., at first convex, soon depressed; surface hygrophanous, dry, subglabrous, at first covered with a minute white soon disappearing pruina (which often fragments concentrically) that persists on the margin, grey avellaneous, tan (5C5–7), brown (4E6–7), dark brown (1F5–7, 3F4–8, 4F6–7), sometimes with purplish-grey tones (7F2–5), margin incurved, uneven, undulate, lobed. Lamellae long decurrent, narrow, very close (crowded) L= 40–70, l= 1–3, some forked halfway, at the base and at the apex, entire, at first white, pale yellowish (3A7–8) when old, with pinkish tones, never turning grey black on handling or bruising, with an even, sinuous, concolorous or paler edge. Stipe 15–30 × 5–8 (–10) mm, usually equal, solitary, often flared at the apex, rarely attenuated at the base, at first white, then beige (5A3), cream avellaneous (6A3–4), white tomentose, especially at the base, with white rhizomorphs at the base. Context very thin, white, unchanging; smell not recorded (of light anise in FLAS-F-71949-iNaturalist-183000711), taste very bitter. Chemical spot-test reactions: KOH on pileus surface negative. Basidiospores (4.3–)4.8– 5.1 –5.4(–6.0) × (3.6–)4.2– 4.5 –4.7(–5.2) µm [192/4/4], Q = (1.00–)1.06– 1.14 –1.21(–1.36), V = (33.3–)45.8– 54.0 –62.3(–75.6) µm 3 , globose, subglobose up to broadly ellipsoid in side and frontal view, clearly angular (9–12 facets) especially when young, obscurely angular when mature, with small pustules or bumps, hilar appendage up to 1 μm long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cianophilic, faintly congophilic, hyaline to pale yellow in water, yellow in Melzer’s (inamyloid). Basidia 18–30(–35) × 6.5–8 μm, cylindrical-clavate, tetrasporic, rarely bisporic, sterigmata up to 5 μm long, usually with guttulate greenish contents. Subhymenium 20–40 μm thick, consisting of textura intricata type short elements, 2–5 μm wide. Hymenophoral trama subregular, hyphae subparallel to intertwined, cylindrical, 2–8 μm wide, thick-walled (wall up to 0.6 µm thick), hyaline to faintly cream. Hymenial cystidia usually absent. In the collection FLAS-F-61088, cystidioid hyaline and thin-walled elements, subcylindrical, sinuous, 20–50 × 2–4 µm, are occasionally present on the lamellar edge (Fig. 7f). Pileipellis as a xerocutis with cylindrical, mostly intertwined hyphae, 2.5–6(–7) μm wide, thick-walled (wall up to 0.6 µm thick), hyaline, smooth, the more superficial ones reclined to ascendant, hyaline or with a pale yellow cytoplasmatic pigment, pileocystidia not observed; thromboplerous hyphae absent. Subpellis of subparallel to loosely intertwined 3–8 μm wide hyaline hyphae. Stipitipellis as a cutis consisting of subparallel to intertwined loose hyphae, 2–5 µm wide, faintly yellowish; caulocystidia as dense tufts of reclining to erect hyphoid elements, often multiseptate, straight to wavy, short to long, 30–100(–150) × 2–4 µm, with an obtuse to acute apex, sometimes also subcapitate, smooth or with an epiparietal pigment in patches. Stipititrama made up of parallel subcylindrical, 3–7 µm wide, hyaline hyphae, thin- to thick-walled (wall up to 0.5 µm thick). Clamp connections absent. Material examined : USA , Florida, Alachua, Prairie Creek Hammock, 29.598463 -82.247989 +-1600 m, on dead leaves, 15 July 1938, leg. West , Arnold & Murrill , FLAS-F-17903 ( Holotype , as Clitocybe alachuana ); Florida, Alachua, Gainesville, 29.66636 -82.32993 +-8000 m, trash in rich hammock, 08 August 1939, leg. E . West , det. WA Murrill , FLAS-F-19879 (as C. alachuana ); Florida, Putnam County, Ordway-Swisher Biological Station, Mill Creek Swamp Bridge, 30.80138 -86.292863, on decorticate wood, Quercus and palm tree dominated swamp, 06 July 2017, leg . D Borland & B Kaminsky , det. ME Smith (as cf. Clitocella ), FLAS-F-61088 ( Epitype ), duplo in CORT014753, det. TJ Baroni (as Lulesia alachuana ); Florida, Melrose, Putnam County, Ordway-Swisher Biological Station, 29.71900833 -81.97166333, on the ground in highly decayed litter and also very decayed wood of hardwoods, 12 September 2023, leg. F Sheffer , FLAS-F-71949-iNaturalist-183000711(https://www.inaturalist.org/observations/183000711) (as Clitocella sp.). Notes – Lulesia alachuana , originally described from Florida (Murrill 1944, Alachua County, inde nomen , from which the species name derives), is so far known only from Florida (Tab. 1 and Fig. 1). The present full description of the species is the first to be provided after the original one by Murrill (1944). Due to its greyish brown pileus, pileipellis as a cutis, subglobose, slightly angular, undulate pustulate or nearly smooth spores, a hymenophoral trama of interwoven hyphae, and bitter taste (Murrill 1944; Bigelow 1982; our observations), L . alachuana is also morphologically a good member of Lulesia subgen. Lulesia , as molecularly supported in the multigene analysis (Fig. 1) where it is sister to a strongly supported clade (BP = 95%, PP = 1.00) consisting of L . mundula , L . obscura and L . popinalis (all with greyish colours, irregular hymenophoral trama and red reaction to KOH). Because of its subglobose spores 5.1 × 4.5 μm, Q = 1.14 on average, and growth on forest (hardwood) litter L. alachuana seems quite close to L . mundula and in fact it was considered a later synonym of the latter by some authors [e.g., T. Baroni’s handwritten notes (1978) accompanying the holotype collection of Clitocybe alachuana (FLAS-F-17903), https://www.mycoportal.org/portal/collections/individual/index.php?occid=604249&clid=0; Baroni (1981); Bigelow (1982, 1985); Singer (1986)]. It morphologically seems to differ from L . mundula mainly by a darker and hygrophanous pileus (brown, dark brown versus white, pale cream, beige-ochraceous), a non-blackening context, a pileus surface with negative KOH reaction (greyish brown versus red) and different nrDNA sequences (Fig. 1; Vizzini et al. 2024 and our observations). Lulesia colorata (L. Fan & N. Mao) T.J. Baroni, N. Niveiro & B.E. Lechner, in Baroni, Lechner & Niveiro, Index Fungorum 566: 1 (2023) Figs. 3g–h, 4b Basionym: Clitocella colorata L. Fan & N. Mao, in Mao, Lv, Xu, Zhao & Fan, MycoKeys 88: 161 (2022) Holotype : BJTC FM1891 For full descriptions (as Clitocella colorata ) see Mao et al. (2022) and Vizzini et al. (2023) Material examined : Finland , Etelä-Karjala, Kouvola, somewhat springfed spruce-hardwood swamp, 19 September 1994, leg. I. Kytövuori (as Rhodocybe cf. mundula ), det. T. Kekki , H6042258 (H). Italy . Lombardia, Pavia, Brallo di Pregola, under Fagus sylvatica , 15 October 2022, leg. M. Carbone , TUR-A 216670; Emilia-Romagna, Piacenza, Ferriere, Loc. Le Moline, mixed forest with Quercus cerris , Q . pubescens and Ostrya carpinifolia , 12 October 2019, leg. et det. M. Carbone & F. Calledda (as Clitocella mundula ), TUR-A 216671. Notes – Lulesia colorata (as Clitocella colorata ) has recently been described from China (Shanxi province, North China or Huabei) on soil or rotten wood in coniferous ( Pinus ) or broad-leaved ( Quercus ) forest (Mao et al. 2022). The Chinese authors highlighted that also a northamerican collection named as C . mundula (TJB7599 (AFTOL-ID 521) USA, NY) clustered within the C . colorata clade. In the original description, no reference is made to a possible colour change of the basidiome surfaces, its smell and taste are not reported, and the hymenophoral trama is indicated as regular (Mao et al. 2022). In their monographic work on the genus Clitocella in Europe, Vizzini et al. (2023), based on molecular data and new collections (from Italy-Veneto and Estonia in mixed coniferous forests) highlighted that C . colorata has: 1) a wider geographical distribution, and it is also present in India, South Korea, USA (Arkansas, Indiana, New York, Tennessee, Wisconsin) and in Europe (France, Italy-Veneto, and Estonia) (Fig. 1); 2) pileus, lamellae, and stipe surface stain strongly black when bruised or in age, smell is farinaceous, taste is bitter, and the hymenophoral trama is subregular at first but then irregular and composed of interwoven hyphae. Our present analysis allow to extend the geographical distribution of this species to Finland and the Italian regions of Lombardy and Emilia-Romagna and to confirm the additional features provided by Vizzini et al. (2023), including also the blackening of basidioma surfaces. Consequently, an emended combination of characters circumscribes L . colorata such as clitocyboid basidiomes (strongly depressed pileus), a usually pale-coloured pileus surface (white to yellowish white, greyish white to greyish brown, pink-white), the surfaces of the basidiome turning black when old or on handling and showing no reddish reaction in KOH, farinaceous smell and bitter taste, (4.7–)5.1– 5.5 –5.9(–6.8) × (3.7–)4.1– 4.3 –4.6(–5.0) µm, (globose) subglobose to broadly ellipsoid basidiospores slightly angular and with minute pustules or bumps, hyphae of pileipellis with pale yellow to yellowish brown intracellular and/or parietal pigment, and the growth in coniferous or angiospermous forests (Mao et al. 2022; Vizzini et al. 2023; our observations). Morphologically, C . colorata can be easily confused with C . mundula and C . popinalis which, however, have dried basidiome surfaces producing a reddish reaction with KOH (Baroni 1981; Moreau 1997; Kluting et al. 2014; Vizzini et al. 2023). Indeed, many collections that clustered in the L . colorata clade (Fig. 1) were originally identified as C . mundula or C . popinalis (Fig. 1 and Tab. 1). Additionally, C. mundula differs in the obscurely angular basidiospores with indistinct pustules or bumps (Baroni 1981; Neukom 1994; Moreau 1997) and C. popinalis in basidiome surfaces usually unchanging on handling, a dark-coloured pileus slightly depressed only in senescing basidiomes, its association with grasses, and slightly broader and longer basidiospores (Baroni 1981; Moreau 1997; Overall 2011; Kluting et al. 2014; Jian et al. 2020a). Lulesia solaris (Musumeci, Consiglio & Vizzini) Musumeci, Consiglio & Vizzini, in Vizzini, Alvarado, Consiglio, Angelini & Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 8. (2024) [2023]. Fig. 4f Basionym: Clitocella solaris Musumeci, Consiglio & Vizzini, in Vizzini et al., Persoonia 50: 144 (2023) Holotype : LUG 19882 For a full description (as C . solaris ) see Vizzini et al. (2023). Material examined : Finland , Kittilän Lappi, Kolari, Pinus sylvestris forest near limestone quarry, 16 August 2001, leg. I. Kytövuori (as Rhodocybe caelata ), det. T. Kekki (H6043012). Notes – The species was recently described from France (Département Haut-Rhin, Alsace) as gregarious on moss pads ( Rhacomitrium canescens ) in open and sunny areas (with seedlings of Betula pubescens and at some distance (at least 30 m) two Pinus sylvestris trees, on sandy-pebbly, alluvial soil, silty substrate rich in woody debris and carbonate). Until the present work it was so far known only from the locus typicus (Vizzini et al. 2023). Lulesia solaris is distinguished by a unique combination of characters such as the small-sized basidiomes (pileus not exceeding 10 mm diam), a whitish cream pileus surface with a positive (red) KOH reaction, distant (spaced) lamellae, an unchanging, not staining black context, subfarinaceous smell and taste, absence of hymenial cystidia and clearly angular, subglobose, broadly ellipsoid to ellipsoid basidiospores with evident bumps in Cotton blue. In the present work,a Finnish twenty-four years old collection misidentified as Rhodocybe caelata (H6043012, see above) turned out to be conspecific with L . solaris (Fig. 1). It was found on calcareous soil and close to Pinus sylvestris trees as the holotype collection. Molecularly, L . solaris is sister (BP = 99 %, PP = 1.00) to a clade formed by C . colorata and C . orientalis (Fig. 1), species which show larger basidiomes, a negative KOH reaction, crowded lamellae, and differently shaped basidiospores (Jian et al. 2020a; Mao et al. 2022; Vizzini et al. 2023 and our observations). Clitocella popinalis has a larger pileus and stipe, a usually dark-coloured pileus surface (purplish grey, brownish grey, blackish grey) crowded lamellae, and subglobose to broadly ellipsoid wider basidiospores (5.1 µm on average) (Vizzini et al. 2023). Discussion The morphological delimited L. fallax species complex is part of Lulesia subgen. Paraclitopilus (pallid Clitopilus -like coloured basidiomes, regular to subregular hymenophoral trama made up of parallel to slightly intertwined cylindrical hyphae, negative KOH reaction, and often adhering in tetrads spores; Vizzini et al. 2023 ). The L. fallax complex consists of species sharing a bitter taste, a white-pruinose pileus reminding Clitocybe sect. Candicantes (Quél.) Singer & Digilio species and often in association with coniferous trees. This species complex encompasses L . fallax , L . neofallax and the two new species L . fallacioides and L . parvifallax , which are all very similar taxa morphologically differentiated almost only based on the size of the basidiomes and the basidiospores (Table 2 ). These species together with those gravitating around L . mundula of Lulesia subgen. Paraclitopilus , viz. L . popinalis , L . solaris , L . colorata , could be considered as cryptic taxa, namely species exhibiting shallow morphological differences (morphological stasis, deceleration of morphological evolution), but considerable genetic disparity (Bickford et al. 2007 ; Grebenc et al. 2009 ; Korhonen et al. 2018 ; Struck et al. 2018a , b ; Korshunova et al. 2019 ; Peintner et al. 2019 ; Cerca et al. 2020 ; Struck and Cerca 2019 , 2022 ; Ekanayaka et al. 2025 ). Cryptic species could be considered as the opposite of adaptive radiation (pronounced morphological differences, shallow genetic divergence) (Cerca et al. 2020 ; Struck and Cerca 2019 , 2022 ) and, a priori, they could result from recent speciation, parallelism, convergence or stasis (Struck and Cerca 2019 , 2022 ). Molecular analyses (Vizzini et al. 2023 , 2024 ; Figs. 1 – 2 in the present paper) would indicate that the phenomena of evolutionary convergence and parallelism should be excluded for the appearance of cryptospecies in Lulesia . Examples of morphological stasis are commonplace in other rhodocyboid fungi, viz. the Rhodophana nitellina complex (Dima et al. 2018 , Buyck et al. 2021 ; Papetti 2023 ), and Rhodocybe gemina complex (Crous et al. 2017 ; Sesli and Vizzini 2017 ; Vizzini et al. 2018 ; Silva-Filho et al. 2020 ; Dutta et al. 2021 ; Sun and Bau 2023 ). As all the species covered in the present paper show difficulty in morphological distinction, these may have been misdetermined in the past, and a greater scrutiny and increased sequencing of historical and modern collections are now imperative before any conclusions can be drawn regarding more precise distribution and rarity data for each of these species in the world. The occurrence of L . fallax in North America (Baroni 1981 ) and The Netherlands (Noordeloos 1988 ) is based on presumably heterogeneous collections and thus giving rise to collective descriptions (basidiospores 6.5–8 × 4–5(–6.5) µm and 5.0–8.5 × 3.5–5.0 µm-Q 1.4–2.0, respectively). So, the presence of the true L . fallax in North America and The Netherlands is still questionable. No sequences in the public databases obtained by Dutch fungal collections clustered within the L . fallax clade and the L . fallax complex (Figs. 1 – 2 ). The only sequences from an American collection named C . fallax (OKM:25668, USA-Oregon), represent the new species L . fallacioides (Fig. 1 ). Two ITS environmental sequences from USA (Utah) (UDB03879566 and UDB03879567) and two basidiomatal sequences (USA-California and China-Xizang) (PP594830 and PV124053) clustered in a clade ( Lulesia sp. 2) within Lulesia subg. Paraclitopilus (Fig. 2 ) which probably represents a distinct not yet described species. Swiss specimens growing on bare soil of a riparian forest named Rhodocybe fallax (Breitenbach & Kränzlin 1995 ), characterized by gracile, diminutive omphaloid basidiomes (pileus 10–20 mm wide, infundibuliform, stipe 10–30 × 1.5–3 mm) with basidiospores 7–8.6 × 3.8–4.8 (Q = 1.7–1.9, V = 76) and resembling Clitopilus sect. Scyphoides Singer (Singer 1986 ; Jian et al. 2020b ), are macromorphologically similar to those cited in Baroni ( 1981 ). They will have to be further studied and molecularly checked to ascertain their taxonomic status. Finally, the presence of L . fallax , L . mundula and L . obscura in Finland is here molecularly confirmed for the first time (Figs. 1 – 2 and see Fig. 4 and Table 1 ). Declarations Acknowledgements Matteo Carbone (Italy), Balint Dima (Hungary), Leanne P. Sheffer, Matthew E. Smith and Caroline B. Willis (USA, Florida), Andrin Gross (Switzerland), Diana Weckman (Finland) are acknowledged for dried fungal material and sequences and photos of some collections. We thank Balint Dima (Hungary), Kare Liimatainen, Tuula Niskanen and Matti Kulju (Finland) for the sequences produced in the FinBOL project and included in this study. Author contribution AV and GC conceived the study, IK, TK and RR conducted sampling and provided materials, MM and RR performed all microscopy analyses mentioned in this study, AV performed the laboratory experiments and prepared all figures and tables, and GC analysed the sequence data. AV wrote the manuscript with contributions from all authors. All authors agreed with the submission of the manuscript. Funding This study has been partly supported by the Finnish Barcode of Life (FinBOL) project which operates under the Finnish Biodiversity Information Facility (FinBIF) Research Infrastructure funded by the Research Council of Finland, and by a grant YM23/5512/2013 from the Finnish Ministry of Environment. The sequencing of Lulesia parvifallax was kindly funded by the Swiss Federal Office for the Environment. Data availability Sequence data have been deposited in GenBank as given in Table 1 and alignments in Figshare (https://doi.org/10.6084/m9.figshare.29654816). Ethics approval and consent to participate Not applicable. Consent for publication Not applicable. Competing interests None References Agerer R (2018) Subphylum Agaricomycotina Doweld. In Begerow D, Agerer R, McTaggart A, Basidiomycota and Entorhizomycota . A. Engler's Syllabus of Plant Families, part 1/3 (Frey W, ed). 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Cite Share Download PDF Status: Published Journal Publication published 21 Nov, 2025 Read the published version in Mycological Progress → Version 1 posted Reviewers agreed at journal 27 Aug, 2025 Reviewers invited by journal 26 Aug, 2025 Editor invited by journal 22 Aug, 2025 Editor assigned by journal 11 Aug, 2025 First submitted to journal 07 Aug, 2025 Editorial decision: Minor Revisions Needed 07 Aug, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7232140","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":506108805,"identity":"08fd1122-2d1e-4aa7-8e79-ebf71b482df6","order_by":0,"name":"Alfredo Vizzini","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA80lEQVRIie3PsWrDMBCA4RMHzuJWq7LYr6CQJUMexl7iqavJZCQE7pIHsN8iY8cUDV5K5kA7uOQF1K0ZCr2EEEhB9tpBPwjd8nESQCj0H5uwHiCiAelkIBIApmgUfoIo78iciCYj/OZGruVqbA03GDlXLkF2k6P7fFkU23fz3DtYVD4iLGLb7FcgLWQyfxNP249XrZuhh1lu8aG2MDWwy/KayCHXJh4gKW3BnwthakekkGNEngkjwulSRLJRMiPCNvtVzDFCIDJribSNFFPlIUlnEL7LZRLx7vh1qqv08VD0zq0r7v3+tfjvg8dAKBQKhYb6BUhTTNrh02ukAAAAAElFTkSuQmCC","orcid":"","institution":"University of Turin: Universita degli Studi di Torino","correspondingAuthor":true,"prefix":"","firstName":"Alfredo","middleName":"","lastName":"Vizzini","suffix":""},{"id":506108806,"identity":"41e6f65b-fd2f-48b3-9329-7e72a8e382b5","order_by":1,"name":"Giovanni Consiglio","email":"","orcid":"","institution":"Independent researcher","correspondingAuthor":false,"prefix":"","firstName":"Giovanni","middleName":"","lastName":"Consiglio","suffix":""},{"id":506108807,"identity":"0bb76de6-bfc0-4d1f-a1e2-e1453c712789","order_by":2,"name":"Tapio Kekki","email":"","orcid":"","institution":"University of Oulu: Oulun Yliopisto","correspondingAuthor":false,"prefix":"","firstName":"Tapio","middleName":"","lastName":"Kekki","suffix":""},{"id":506108808,"identity":"d0c7676a-e7ce-4eee-bcb9-ce1b7675f5b3","order_by":3,"name":"Mauro Marchetti","email":"","orcid":"","institution":"Independent researcher","correspondingAuthor":false,"prefix":"","firstName":"Mauro","middleName":"","lastName":"Marchetti","suffix":""},{"id":506108809,"identity":"0f9a8495-75ec-4802-93f5-1112fd8de523","order_by":4,"name":"Ilkka Kytövuori","email":"","orcid":"","institution":"Finnish Museum of Natural History: Luonnontieteellinen museo","correspondingAuthor":false,"prefix":"","firstName":"Ilkka","middleName":"","lastName":"Kytövuori","suffix":""},{"id":506108810,"identity":"d86b327c-e2ef-4626-a7b3-1a099e411996","order_by":5,"name":"Raphael Rickmann","email":"","orcid":"","institution":"Independent researcher","correspondingAuthor":false,"prefix":"","firstName":"Raphael","middleName":"","lastName":"Rickmann","suffix":""}],"badges":[],"createdAt":"2025-07-28 09:28:01","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7232140/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7232140/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11557-025-02100-4","type":"published","date":"2025-11-21T15:57:07+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":90524242,"identity":"b8b77e1a-8ff1-442f-9567-d4be6c79bd26","added_by":"auto","created_at":"2025-09-03 16:31:57","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":343129,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum likelihood\u003cstrong\u003e \u003c/strong\u003ephylogram built with nucleotide sequence data of the nrITS/nrLSU/ \u003cem\u003eRPB2\u003c/em\u003e/\u003cem\u003eTEF-1α\u003c/em\u003e/\u003cem\u003eATP6\u003c/em\u003e dataset of \u003cem\u003eLulesia\u003c/em\u003e, rooted with \u003cem\u003eClitopilus prunulus \u003c/em\u003e(\u003cem\u003eEntolomataceae\u003c/em\u003e). Nodes were annotated on or below the branches with BP ≥ 70 % (left) and PP ≥ 0.95 (right). Subsignificant support values were annotated in parentheses. Newly sequenced collections are in bold. The lengths of the branches were estimated as average values on the sampled trees.\u003c/p\u003e","description":"","filename":"FIG1dapubblicare.png","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/10588fe52ab464858629d076.png"},{"id":90524241,"identity":"5e0d2da2-7db9-4e1e-b809-24e43c87881b","added_by":"auto","created_at":"2025-09-03 16:31:57","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":74324,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum likelihood\u003cstrong\u003e \u003c/strong\u003ephylogram built with nucleotide sequence data of the nrITS dataset of \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e, rooted with \u003cem\u003eL\u003c/em\u003e. \u003cem\u003epopinalis \u003c/em\u003e(\u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eLulesia\u003c/em\u003e). Nodes were annotated on or below the branches with BP ≥ 70 % (left) and PP ≥ 0.95 (right). Subsignificant support values were annotated in parentheses. Newly sequenced collections are in bold.\u003c/p\u003e","description":"","filename":"FIG2.png","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/cc825e3da93257af1dcef5ec.png"},{"id":90524004,"identity":"09c8234e-c35d-40be-9880-fef508d04463","added_by":"auto","created_at":"2025-09-03 16:23:58","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":40924850,"visible":true,"origin":"","legend":"\u003cp\u003eFresh\u003cstrong\u003e \u003c/strong\u003ebasidiomes of sequenced collections. \u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003ec\u003c/strong\u003e \u003cem\u003eLulesia fallacioides \u003c/em\u003e[OULU:GAJ.17963-T. Kekki (6650), OULU:GAJ.17962-T. Kekki (6670) Holotype, OULU:GAJ.11332-T. Kekki (3988)]. \u003cstrong\u003ed\u003c/strong\u003e–\u003cstrong\u003ee\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax \u003c/em\u003e(G00586348, Holotype). \u003cstrong\u003ef\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula \u003c/em\u003e(OULU:GAJ.17966). \u003cstrong\u003eg\u003c/strong\u003e–\u003cstrong\u003eh\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata \u003c/em\u003e(TUR-A 216670, TUR-A 216671). \u003cstrong\u003ei\u003c/strong\u003e–\u003cstrong\u003ej\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ealachuana \u003c/em\u003e(FLAS-F-71949-iNaturalist-183000711). Photos: a–c, f, by T. Kekki; d–e by R. Rickmann; g–h by M. Carbone; i–j by L.P. Sheffer.\u003c/p\u003e","description":"","filename":"FIG3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/734e925f947c9e62b706b310.jpg"},{"id":90523995,"identity":"a54a2be1-8e93-4fea-89aa-fb412beaecbf","added_by":"auto","created_at":"2025-09-03 16:23:57","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":19299061,"visible":true,"origin":"","legend":"\u003cp\u003eDried\u003cstrong\u003e \u003c/strong\u003ebasidiomes of sequenced collections. \u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003eb \u003c/strong\u003e\u003cem\u003eLulesia fallacioides\u003c/em\u003e (H6032673, H6032677). \u003cstrong\u003ec\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e(H6032675). \u003cstrong\u003ed\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata \u003c/em\u003e(H6042258). \u003cstrong\u003ee\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e (H6042309)\u003cstrong\u003e f\u003c/strong\u003e–\u003cstrong\u003eg \u003c/strong\u003e\u003cem\u003eL\u003c/em\u003e. \u003cem\u003eobscura \u003c/em\u003e(H6042319, H6042347). \u003cstrong\u003eh\u003c/strong\u003e \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e (H6043012). Photos: a–c by D. Weckman; d–h by B. Dima.\u003c/p\u003e","description":"","filename":"FIG4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/6734d32756cf5033cf6af3d1.jpg"},{"id":90523991,"identity":"955ac0ad-00bf-4fd2-99e1-097eb1961f66","added_by":"auto","created_at":"2025-09-03 16:23:57","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":12325116,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eLulesia fallacioides\u003c/em\u003e. Microscopic characters. \u003cstrong\u003ea\u003c/strong\u003e Pileipellis. \u003cstrong\u003eb\u003c/strong\u003e Hymenium (basidia). \u003cstrong\u003ec\u003c/strong\u003eCaulocystidia. \u003cstrong\u003ed\u003c/strong\u003e–\u003cstrong\u003eh\u003c/strong\u003e Basidiospores. a–f from OULU:GAJ.17962-T. Kekki (6670) Holotype, g from OULU:GAJ.17963-T. Kekki (6650), h from OULU:GAJ.11332-T. Kekki (3988). a in phloxine, b–e, g–h in Congo red, f in Cotton blue. a–c, e–f Bright-field microscopy, d, g–h interference contrast microscopy. Photos by M. Marchetti. Scale bars: a = 30 μm, b–h = 10 μm. Photos by M. Marchetti.\u003c/p\u003e","description":"","filename":"FIG5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/574a91aacf577754da379e63.jpg"},{"id":90523992,"identity":"b3ef8aab-194c-4608-b80b-b9ef63fbde28","added_by":"auto","created_at":"2025-09-03 16:23:57","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":7719816,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eLulesia parvifallax\u003c/em\u003e. Microscopic characters (G00586348, Holotype). \u003cstrong\u003ea\u003c/strong\u003e Basidiospores. \u003cstrong\u003eb\u003c/strong\u003eBasidia. \u003cstrong\u003ec \u003c/strong\u003ePileipellis \u003cstrong\u003ed\u003c/strong\u003e Elements of the hymenophoral trama. Scale bars = 10 μm. Drawings by R. Rickmann.\u003c/p\u003e","description":"","filename":"FIG6oKconlettere.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/11451d0b7cee92d8954b71a0.jpg"},{"id":90524001,"identity":"6b5905c3-cff5-439a-baf4-32df1acda5ec","added_by":"auto","created_at":"2025-09-03 16:23:58","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":24713712,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eLulesia alachuana\u003c/em\u003e. \u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003eb\u003c/strong\u003ePileipellis (FLAS-F-71949-iNaturalist-183000711, FLAS-F-61088, Epitype). \u003cstrong\u003ec\u003c/strong\u003ePileipellis scalp (FLAS-F-71949-iNaturalist-183000711). \u003cstrong\u003ed\u003c/strong\u003e–\u003cstrong\u003ee\u003c/strong\u003eHymenophoral trama (FLAS-F-71949-iNaturalist-183000711, FLAS-F-61088, Epitype). \u003cstrong\u003ef \u003c/strong\u003eCystidioid (rare) hyphoid elements on the lamellar edge (FLAS-F-61088, Epitype). \u003cstrong\u003eg\u003c/strong\u003e–\u003cstrong\u003eh\u003c/strong\u003e Caulocystidia (FLAS-F-61088, Epitype, FLAS-F-71949-iNaturalist-183000711). a–c, e–h in Congo red, d in Cotton blue. a–g Bright-field microscopy, h interference contrast microscopy. Scale bars: a = 20 μm, b–f, h = 10 μm, g = 30 μm. Photos by M. Marchetti.\u003c/p\u003e","description":"","filename":"FIG7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/d47e92039b3b4db64a286b18.jpg"},{"id":90523993,"identity":"dacfe0fc-a6e1-4d0d-9c78-339584b32bdb","added_by":"auto","created_at":"2025-09-03 16:23:57","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":6666716,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eLulesia alachuana\u003c/em\u003e. \u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003eb\u003c/strong\u003eHymenium (basidia and developing spores) (FLAS-F-61088, Epitype, FLAS-F-71949-iNaturalist-183000711). \u003cstrong\u003ec\u003c/strong\u003e–\u003cstrong\u003ef\u003c/strong\u003e Basidiospores (FLAS-F-19879, FLAS-F-71949-iNaturalist-183000711, FLAS-F-71949-iNaturalist-183000711, FLAS-F-61088, Epitype). a–d, f in Congo red, e in Cotton blue. Scale bars = 10 μm. All photos in interference contrast microscopy. Photos by M. Marchetti.\u003c/p\u003e","description":"","filename":"FIG8.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/9ba3fbae2886eae9aaf5d314.jpg"},{"id":96651414,"identity":"5a3aec9b-f6a2-468c-a9fb-d477b9dcd549","added_by":"auto","created_at":"2025-11-24 16:14:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":115177706,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7232140/v1/6f09ce6e-1bc3-4b4d-9db4-4402263f9c92.pdf"}],"financialInterests":"","formattedTitle":"Two new species in the Lulesia fallax complex (Entolomataceae, Agaricales) from Europe (Fennoscandia and Switzerland), Lulesia neofallax comb. nov., new records of recently described species of Lulesia, and notes on Clitocybe alachuana","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe genus \u003cem\u003eLulesia\u003c/em\u003e was established by Singer (\u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e1970\u003c/span\u003e) for accommodating two species previously considered as part of \u003cem\u003eArmillariella\u003c/em\u003e Singer (Singer \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e1951\u003c/span\u003e; Singer and Digilio \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e1951\u003c/span\u003e), viz. \u003cem\u003eA\u003c/em\u003e. \u003cem\u003edensifolia\u003c/em\u003e Singer (type of \u003cem\u003eLulesia\u003c/em\u003e, from Tucum\u0026aacute;n, Argentina) and \u003cem\u003eClitocybe alachuana\u003c/em\u003e Murrill (from Florida, USA). Singer (\u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e1970\u003c/span\u003e) differentiated \u003cem\u003eLulesia\u003c/em\u003e from \u003cem\u003eArmillariella\u003c/em\u003e by having a zonate pileus, very narrow lamellae, a bitter taste, smaller spores, a trichodermic pileus covering, and a terricolous/humicolous habitat. \u003cem\u003eLulesia densifolia\u003c/em\u003e spores were described as smooth in some mounting media (methylene blue, carmine-acetic acid, and Melzer\u0026rsquo;s reagent) but appearing slightly rounded-angular and nodulose in aqueous NH\u003csub\u003e3\u003c/sub\u003e and KOH in light microscopy (Singer and Digilio \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e1951\u003c/span\u003e; Singer \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e1970\u003c/span\u003e, \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). According to Singer and Digilio (\u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e1951\u003c/span\u003e) and Singer (\u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e1986\u003c/span\u003e) this slight irregularity of the spore surface would be reminiscent of the spores of the species in genus \u003cem\u003eRhodocybe\u003c/em\u003e Maire.\u003c/p\u003e\u003cp\u003eBaroni (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1981\u003c/span\u003e) and Bigelow (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1982\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1985\u003c/span\u003e), after studying the holotype collection of \u003cem\u003eClitocybe alachuana\u003c/em\u003e, considered it a later synonym of \u003cem\u003eRhodocybe mundula\u003c/em\u003e (Lasch) Singer, and this conclusion was also followed by Singer (\u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). The species recognized in \u003cem\u003eLulesia\u003c/em\u003e were recently supplemented by Lechner et al. (\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) with another new species from Argentina, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003elignicola\u003c/em\u003e B.E. Lechner \u0026amp; J.E. Wright. No further collections of \u003cem\u003eL. densifolia\u003c/em\u003e apart from the original ones were reported until recent findings in the Dominican Republic (Angelini and Contu \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eBased only on morphological data, \u003cem\u003eLulesia\u003c/em\u003e was placed by Singer (\u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e1970\u003c/span\u003e, \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e1986\u003c/span\u003e) in subtribe \u003cem\u003eOmphalineae\u003c/em\u003e Singer (tribe \u003cem\u003eClitocybeae\u003c/em\u003e Fayod, family \u003cem\u003eTricholomataceae\u003c/em\u003e R. Heim ex Pouzar), by Lechner et al. (\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) and Agerer (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) in \u003cem\u003eTricholomataceae\u003c/em\u003e s.l., and by He et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) in \u003cem\u003eincertae sedis Agaricales\u003c/em\u003e Underw. and \u003cem\u003eEntolomataceae\u003c/em\u003e Kotl. \u0026amp; Pouzar, respectively.\u003c/p\u003e\u003cp\u003eRecently, based on multigene analyses, Vizzini et al. (\u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) have highlighted that \u003cem\u003eL. densifolia\u003c/em\u003e clustered within the genus \u003cem\u003eClitocella\u003c/em\u003e Kluting, T.J. Baroni \u0026amp; Bergemann, a genus (typified by \u003cem\u003eAgaricus popinalis\u003c/em\u003e Fr.) segregated from \u003cem\u003eRhodocybe\u003c/em\u003e s.l. (\u003cem\u003eEntolomataceae\u003c/em\u003e) (Kluting et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and recently monographed at European level by Vizzini et al. (\u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This indicated \u003cem\u003eLulesia\u003c/em\u003e as a priority synonym of \u003cem\u003eClitocella\u003c/em\u003e. Three distinct subgenera are currently recognized in \u003cem\u003eLulesia\u003c/em\u003e relied on type of hymenophoral trama, basidiome colours and reactions of basidiome surface to KOH: subgen. \u003cem\u003eLulesia\u003c/em\u003e, \u003cem\u003eParaclitopilus\u003c/em\u003e (Vizzini \u0026amp; Consiglio) Vizzini \u0026amp; Consiglio, and \u003cem\u003eRhodopleurella\u003c/em\u003e (Vizzini \u0026amp; Consiglio) Vizzini \u0026amp; Consiglio (Vizzini et al. \u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cem\u003eLulesia\u003c/em\u003e species typically produce a gymnocarpic centrally stipitate to pleuropodal (\u003cem\u003eL\u003c/em\u003e. \u003cem\u003etermitophila\u003c/em\u003e (T.J. Baroni \u0026amp; Angelini) T.J. Baroni \u0026amp; Angelini), clitocyboid basidiome, which exhibits a small to large (\u0026le;\u0026thinsp;120 mm) pileus that can range from white to greyish, grey-brown or purplish grey in colour, usually covered with a white bloom, long-decurrent narrow lamellae with pinkish tinges at maturity, a cylindrical glabrous to pubescent, floccose or fibrillose stipe usually with white rhizomorphs at the base, context usually with farinaceous to cucumber-like smell and taste bitter, surfaces of the dried basidiomes reddening or not in KOH, spore-print incarnate-pink, hymenophoral trama regular (consisting of parallel cylindrical hyphae) or irregular (composed by interwoven hyphae), hymenial cystidia usually lacking, and cyanophilic, inamyloid, non-dextrinoid, thin-walled basidiospores with obscure or distinct undulating pustules or minute bumps (Kluting et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e, \u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). To date, a total of 17 species of \u003cem\u003eLulesia\u003c/em\u003e and two uncombined \u003cem\u003eClitocella\u003c/em\u003e species have been validly published based on the records from the Index Fungorum database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.indexfungorum.org\" target=\"_blank\"\u003ewww.indexfungorum.org\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.indexfungorum.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, accessed 21 January 2025). \u003cem\u003eLulesia\u003c/em\u003e has been reported in Asia, Europe, North, South and Central America, and can be observed in various ecosystems, including forested and grassland systems, as well as on dunes or in association with hind dune trees, from temperate to subtropical/tropical areas (Angelini and Contu \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kluting et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Baroni et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Jian et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e; Mao et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e, \u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Xiao et al. \u003cspan citationid=\"CR149\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Its species are presumably saprotrophic (Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e, \u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), usually terrestrial but \u003cem\u003eC\u003c/em\u003e. \u003cem\u003elignicola\u003c/em\u003e is reported on dead wood of dicotyledonous trees (Lechner et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e)d \u003cem\u003etermitophila\u003c/em\u003e grows on arboreal nests of neotropical termites (Baroni et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe aim of the present work was to describe two new white \u003cem\u003eLulesia\u003c/em\u003e species from Europe morphologically closely related to \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, by using both a traditional (morphological) and multilocus molecular approach. Additionally, a full morphological description of the north American species \u003cem\u003eClitocybe alachuana\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eL\u003c/em\u003e. \u003cem\u003ealachuana\u003c/em\u003e (Murrill) Singer) based on ancient and modern collections, and notes on the recently established species \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e (Mao et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), are provided.\u003c/p\u003e"},{"header":"Materials \u0026 Methods","content":"\u003cp\u003e\u003cem\u003e\u003cu\u003eMorphological studies\u003c/u\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eMacroscopic morphological features were studied in fresh specimens. Codes for colours follow the Methuen Handbook of Colour (Kornerup and Wanscher 1984). The following abbreviations are employed: L = number of lamellae reaching the stipe; l = number of lamellulae between each pair of lamellae. Microscopic structures were examined in dried material using different mounting media: water, L4 (Cl\u0026eacute;men\u0026ccedil;on 1972), Melzer\u0026rsquo;s reagent, ammoniacal Congo red, phloxine, and Cotton blue. Dried pieces of the samples were rehydrated in water and mounted in L4. All microscopic measurements were carried out with a Nikon Eclipse 80i microscope, using immersion oil at \u0026times;1000 magnification. Basidiospore measurements were made by capturing images of a single visual field with multiple spores (taken from lamellar squashes of exsiccated material of mature specimens) which were then measured using the DS-L1 Nikon camera control unit. Basidiospore dimensions excluded the hilar appendix and are given as (a)b\u0026ndash;\u003cem\u003ec\u003c/em\u003e\u0026ndash;d(e), where (a) = minimum value, b = (average minus standard deviation), c = \u003cem\u003eaverage\u003c/em\u003e, d = (average plus standard deviation) and (e) = maximum value; Q = (minimum\u0026minus;) average minus standard deviation \u0026ndash; \u003cem\u003eaverage\u003c/em\u003e \u0026ndash; average plus standard deviation (\u0026minus;maximum) of ratio length/width; V = (minimum\u0026minus;) average minus standard deviation \u0026ndash; \u003cem\u003eaverage\u003c/em\u003e \u0026ndash; average plus standard deviation (\u0026minus;maximum) of the volume. The approximate spore volume was calculated as that of an ellipsoid (Gross 1972; Meerts 1999). The notation [n/m/p] indicates that measurements were made on \u0026lsquo;n\u0026rsquo; randomly selected spores from \u0026lsquo;m\u0026rsquo; basidiomes of \u0026lsquo;p\u0026rsquo; collections. A minimum of 30 spores were measured for each collection. Q indicates the quotient of length and width of the spores in side view. The width of the basidia was measured at the widest part, and the length was measured from the apex (sterigmata excluded) to the basal septum. Photographs of microscopic features were taken using a Nikon DS 5M digital camera with a preset resolution of 2560 \u0026times; 1980 pixels connected to a Nikon Eclipse 80i microscope with both brightfield and interferential contrast optics and saved in TIFF format. They were then cleaned using Adobe Photoshop CC 2019 version 20.0.10, in some cases replacing, through the Magic Wand Tool and with the appropriate tolerances, the background colour of the photograph with an alternative uniform background to highlight the morphology of the microscopic characters, which maintained their original size and hue. In other cases, the best individual spores from microscopy photos were copied using the Polygonal Lasso Tool or alternatively the Magnetic Lasso Tool of Adobe Photoshop CC 2019 version 20.0.10, onto a new monochrome sheet, with the same resolution (2560 \u0026times; 1980 pixels) to maintain the spore size and original colours as previously acquired.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eMacro- and microchemical testing of pigments were performed using basic solutions (5% KOH and 10% ammonia, separately). Chemical spot tests were performed on pileus surface, lamellae and stipe of fresh and/or dried basidiomes using 3% and 10% KOH, following Baroni (1978, 1981). Fungarium acronyms follow Thiers (2025) except that ANGE and T. Kekki refer to the personal herbarium of Claudio Angelini and Tapio Kekki, respectively.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u003cu\u003eDNA extraction, amplification, and sequencing\u003c/u\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTotal DNA was extracted from seventeen dry specimens (Fig. 1, Tab. 1) employing a modified protocol based on Murray and Thompson (1980). PCR (Mullis and Faloona 1987) reaction conditions (cycles and annealing temperature) follow Alvarado (2010, 2012). The primers ITS1F and ITS4 (White et al. 1990; Gardes and Bruns 1993) were used to amplify the ITS rDNA nuclear region, LR0R and LR5 (Vilgalys and Hester 1990; Cubeta et al. 1991) for the LSU rDNA nuclear region, EF1-983F, EF1-1567R and EF1- 2218R (Rehner and Buckley 2005) for the translation elongation factor 1-alpha (\u003cem\u003eTEF1-1\u0026alpha;\u003c/em\u003e) gene, and bRPB2-6F2 (reverse of bRPB2-6R2), and bRPB2-7R2 for the RNA polymerase II second largest subunit (\u003cem\u003eRPB2\u003c/em\u003e) gene (Matheny et al. 2007). PCR products were checked in 1% agarose gels, and amplicons were sequenced with one or both PCR primers. Sequences were corrected to remove reading errors in chromatograms.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u003cu\u003ePhylogenetic analyses\u003c/u\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTwo different dataset/alignments were built. 1) A \u003cem\u003eLulesia\u003c/em\u003e-dataset aimed to provide an accurate view of the position of the newly sequenced collections within this genus was prepared. It included sequences of five different loci (nrITS, nrLSU, \u003cem\u003eRPB2\u003c/em\u003e-exons, \u003cem\u003eTEF-1\u0026alpha;\u003c/em\u003e -exons and \u003cem\u003eATP6\u003c/em\u003e) from all the species recognized in Vizzini et al. (2023, 2024) supplemented with those present in Xiao et al. (2024) and Liu et al. (2025), and the most closely related sequences selected from public databases as the International Nucleotide Sequence Database Collaboration public database (INSDC/GenBank https://www.ncbi.nlm.nih. gov/genbank/,\u0026nbsp;Arita et al. 2021), UNITE (https://unite.ut.ee/), and BOLDSYSTEMS (http://www.boldsystems.org/), via BLASTn algorithm (Altschul et al. 1990). \u003cem\u003eClitopilus prunulus\u003c/em\u003e was employed as outgroup taxon following\u0026nbsp;Baroni\u0026nbsp;et al. (2020), Mao et al. (2022) and Vizzini et al. (2023, 2024). 2) A nrITS dataset focused on \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e and including also all environmental sequences was prepared, with \u003cem\u003eC\u003c/em\u003e. \u003cem\u003epopinalis\u003c/em\u003e as\u0026nbsp;outgroup taxon. All the sequences employed are listed in Table 1.\u0026nbsp;The combined and ITS alignments used for the phylogenetic analyses are deposited in Figshare (https://doi.org/10.6084/m9.figshare.29654816).\u003c/p\u003e\n\u003cp\u003eSequences first were aligned in MEGA 6.0 (Tamura et al. 2013) with its MUSCLE (Edgar 2004) applications and then realigned manually as needed to establish positional homology.\u0026nbsp;The matrices were analysed by using the Bayesian Inference (BI) and the Maximum Likelihood (ML) criteria through the MESQUITE 3.81 (Maddison and Maddison 2023) software by which were obtained the .nex and .phy files.\u003c/p\u003e\n\u003cp\u003eThe .nex files were loaded into MrBayes v. 3.2.7a (Ronquist et al. 2012) of the CIPRES Science Gateway v. 3.3 platform (Miller et al. 2010) and Bayesian analyses were performed with the nrITS\u0026ndash;nrLSU\u0026ndash;\u003cem\u003eRPB2\u003c/em\u003e\u0026ndash;\u003cem\u003eTEF-1\u003c/em\u003e\u003cem\u003e\u0026alpha;\u003c/em\u003e-\u003cem\u003eATP6\u003c/em\u003e partitioned alignment and the ITS alignment (GTR+G+I model, two simultaneous runs, six chains, temperature set at 0.2, sampling every 1 000 generations) until reaching convergence parameters (standard deviation less than 0.01 and PSRF (Potential Scale Reduction Factor) (Gelman and Rubin 1992) equal to 1), after 3.17 M generations and 13.45 M generations, respectively. As required from the procedure, 25 % of the trees, those of the initial stretch and those of the\u0026nbsp;final tail, were \u0026lsquo;burned\u0026rsquo;.\u003c/p\u003e\n\u003cp\u003eA full search for the best-scoring maximum likelihood tree was performed loading the .phy files of the multilocus alignment and ITS alignment into the RAxML v. 8.2.12 \u0026nbsp;program (Stamatakis 2014) using the standard search algorithm with same partitions and 1 000 bootstrap cycles according to the GTR + GAMMA model. The trees in .tre format were read with the software SEAVIEW v. 4 (Gouy et al. 2010) and saved in a vector format for printing.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe significance threshold was set \u0026ge; 70% for bootstrap proportions (BP) and \u0026ge; 0.95 for posterior probabilities (PP).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003e\u003cu\u003eMolecular data\u003c/u\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe topology of the best tree from the ML analysis and 50 % majority-rule consensus tree from the BI analysis are essentially identical, so only the ML tree is presented here, with support values from both analyses (Bootstrap (BS) \u0026ge; 70 % and/or Posterior Probabilities (PP) \u0026ge; 0.95) (Figs 1\u0026ndash;2).\u003c/p\u003e\n\u003cp\u003eThe final combined multigene nrITS\u0026ndash;nrLSU\u0026ndash;\u003cem\u003eRPB2\u003c/em\u003e\u0026ndash;\u003cem\u003eTEF-1\u0026alpha;\u003c/em\u003e\u0026ndash;\u003cem\u003eATP6\u003c/em\u003e \u003cem\u003eLulesia\u003c/em\u003e data matrix encompassed a total of 339 sequences (including 27 newly generated, 303 from INSDC/GenBank and 9 from UNITE) (100 ITS, 66 LSU, 84 \u003cem\u003eRPB2\u003c/em\u003e, 51 \u003cem\u003eTEF-1\u0026alpha;\u003c/em\u003e, 38 \u003cem\u003eATP6\u003c/em\u003e) from 132 samples of 21 taxa. The alignment is 3387 bp long (gaps included).\u003c/p\u003e\n\u003cp\u003eThe nrITS \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e data matrix encompassed a total of 34 sequences (including 7 newly generated, 22 from INSDC/GenBank and 5 from UNITE) from 34 samples of 8 taxa. The alignment is 604 bp long (gaps included).\u003c/p\u003e\n\u003cp\u003eExcept for \u003cem\u003eL\u003c/em\u003e. \u003cem\u003etermitophila\u003c/em\u003e, all the other species of \u003cem\u003eLulesia\u003c/em\u003e included in the present multigene analysis clustered together forming a monophyletic lineage (BP = 100%, PP = 1.00). \u003cem\u003eLulesia termitophila\u003c/em\u003e was sister to all the other species of \u003cem\u003eLulesia\u003c/em\u003e (BP = 100%, PP = 1.00) (Fig. 1). Three major clades are recognized in \u003cem\u003eLulesia\u003c/em\u003e, corresponding to \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eLulesia\u0026nbsp;\u003c/em\u003e(BP = 92%, PP = 1.0), subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e (BP = 92%, PP = 1.00) and subgen. \u003cem\u003eRhodopleurella\u003c/em\u003e, as previously highlighted by Vizzini et al. (2023, 2024).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eLulesia\u003c/em\u003e encompasses nineteen monophyletic species-rank clades corresponding to fifteen already described species, two new species and two \u003cem\u003eLulesia\u003c/em\u003e sp. (\u003cem\u003eLulesia\u003c/em\u003e sp. 1 and \u003cem\u003eLulesia\u003c/em\u003e sp. 2) (Fig. 1). All these species clades with more than one sequence were strongly supported. The two new species (described here as \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e) and \u003cem\u003eLulesia\u003c/em\u003e sp. 2 belong to \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u0026nbsp;\u003c/em\u003ewhich also includes \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eblancii\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eneofallax\u003c/em\u003e (Fig. 1). \u003cem\u003eLulesia\u003c/em\u003e \u003cem\u003efallacioides\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u0026nbsp;\u003c/em\u003eand \u003cem\u003eLulesia\u003c/em\u003e sp. 2 (as \u003cem\u003eLulesia\u003c/em\u003e sp.) are clearly supported as distinct taxa also in the nrITS analysis focused on \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u0026nbsp;\u003c/em\u003e(Fig. 2).\u003cem\u003e\u0026nbsp;Lulesia\u003c/em\u003e sp. 2 consists of two environmental sequences from USA (Utah) and two basidiomatal sequences (China-Xizang and USA-California). Morphological data will be needed to formally describe the clade as a new species.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u003cu\u003eTaxonomy\u003c/u\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003esubgen.\u003cem\u003e\u0026nbsp;Paraclitopilus\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e(Vizzini \u0026amp; Consiglio) Vizzini \u0026amp; Consiglio, in Vizzini, Alvarado, Consiglio, Angelini \u0026amp; Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 8 (2024) [2023]\u003c/p\u003e\n\u003cp\u003eBasionym: \u003cem\u003eClitocella\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e Vizzini \u0026amp; Consiglio, Persoonia 50: 146 (2023)\u003c/p\u003e\n\u003cp\u003eType: \u003cem\u003eOmphalia fallax\u003c/em\u003e Qu\u0026eacute;l., C. r. Assoc. Fran\u0026ccedil;. Avancem. Sci. 24(2): 617 (1896) [1895]\u003c/p\u003e\n\u003cp\u003eHabit like that of the centrally-stipitate \u003cem\u003eClitopilus\u003c/em\u003e species, pileus white to cream coloured under a white pruinose covering, surfaces of both fresh and dried basidiomes not staining grey or black when bruised or in age and with a negative KOH reaction, hymenophoral trama regular, basidiospores broadly ellipsoid, ellipsoid to oblong, amygdaliform, obscurely angular, often adhering (grouping) in tetrads, and sporal Q on average usually exceeding (1.20\u0026ndash;)1.30.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia fallacioides\u003c/em\u003e\u003c/strong\u003eKekki, Vizzini, Consiglio, M. Marchetti \u0026amp; Kyt\u0026ouml;v., \u003cstrong\u003e\u003cem\u003esp. nov.\u0026nbsp;\u003c/em\u003e\u003c/strong\u003eFigs. 3a\u0026ndash;c, 5\u003c/p\u003e\n\u003cp\u003eMycoBank MBXXXXXXX\u003c/p\u003e\n\u003cp\u003eEtymology: fallacio\u0026igrave;des = from the root fallac- of fallax (fallac-is) (Latin) and the suffix -\u0026igrave;des relating to Greek \u0026epsilon;\u0026iota;\u0026delta;\u0026omicron;\u0026sigmaf; (\u0026eacute;idos), which means \u0026ldquo;external aspect, form, semblance\u0026rdquo;, therefore indicating resemblance to \u003cem\u003eLulesia\u003c/em\u003e \u003cem\u003efallax\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDiagnosis\u003c/em\u003e:\u003cem\u003e\u0026nbsp;\u003c/em\u003eIt differs from \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u0026nbsp;\u003c/em\u003eby an ixocutis type of pileipellis, wider basidiospores (4.7 \u0026mu;m on average), and association with \u003cem\u003ePicea abies\u003c/em\u003e forests \u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHolotype\u003c/em\u003e (here designated): OULU:GAJ.17962\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePileus\u003c/em\u003e 10\u0026ndash;50(\u0026ndash;80) mm diam., convex, then applanate and usually depressed in the centre, often irregular when old. Margin involute when young, then straight, usually strongly wavy and undulating; surface dry, smooth to slightly tomentose, not hygrophanous, not translucently striate, white, sometimes with slight cream or ochre tinges in the centre when old. \u003cem\u003eLamellae\u003c/em\u003e L = 20\u0026ndash;45, l = (2\u0026ndash;) 3\u0026ndash;5, crowded, arcuate-decurrent, 2\u0026ndash;4 mm broad, sordid white, finally tinged with pink, pinkish puff to clay-puff tinges when dry, with a smooth concolorous edge. \u003cem\u003eStipe\u003c/em\u003e 8\u0026ndash;26 \u0026times; 2\u0026ndash;7 mm, cylindrical, often tapering downwards, subclavate at the base, solid, white, usually with white rhizomorphs at the base. \u003cem\u003eContex\u0026nbsp;\u003c/em\u003esolid, white. \u003cem\u003eSmell\u003c/em\u003e not distinctive, \u003cem\u003etaste\u003c/em\u003e bitter. No part of the basidiome staining black. \u003cem\u003eSpore-print\u003c/em\u003e pale cream pink. \u003cem\u003eChemical spot-test reactions\u003c/em\u003e KOH on dried basidiomes surfaces (pileus, lamellae and stipe) produces no reaction (negative).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eBasidiospores\u003c/em\u003e(4.81\u0026ndash;)5.9\u0026ndash;\u003cem\u003e6.5\u003c/em\u003e\u0026ndash;7.2(\u0026ndash;8.45) \u0026times; (3.59\u0026ndash;)4.4\u0026ndash;\u003cem\u003e4.7\u003c/em\u003e\u0026ndash;5.1(\u0026ndash;5.70) \u0026micro;m [175/3/3], Q = (1.10\u0026ndash;)1.27\u0026ndash;\u003cem\u003e1.38\u003c/em\u003e\u0026ndash;1.50(\u0026ndash;1.79), V = (37.9\u0026ndash;)59.2\u0026ndash;\u003cem\u003e76.8\u003c/em\u003e\u0026ndash;94.5(\u0026ndash;138) \u0026mu;m\u003csup\u003e3\u003c/sup\u003e\u003cstrong\u003e,\u0026nbsp;\u003c/strong\u003esubamygdaliform in side view, ovoid-ellipsoid in frontal view, minutely angular in polar view (7\u0026ndash;10 facets), weakly nodulose-pustulate to verrucose in all views, rounded to slightly attenuated apically, often adhering in tetrads, thin-walled (no sclerospores), hilar appendage evident, up to 1 \u0026mu;m long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cyanophilic, pale yellow in water, yellow in Melzer\u0026rsquo;s. \u003cem\u003eBasidia\u003c/em\u003e 18\u0026ndash;30 \u0026times; 7\u0026ndash;10 \u0026mu;m, cylindrical to clavate-cylindrical, usually with guttulate contents, mainly tetrasporic, rarely bisporic, sterigmata up to 4 \u0026mu;m long. \u003cem\u003eSubhymenium\u003c/em\u003e 20\u0026ndash;30 \u0026mu;m thick, consisting of \u003cem\u003etextura intricata\u003c/em\u003e short elements, 2.5\u0026ndash;5 \u0026mu;m wide. \u003cem\u003eHymenophoral trama\u003c/em\u003e regular to subregular of parallel to slightly intervowen cylindrical, thin-walled hyphae, hyaline, 3\u0026ndash;10 \u0026mu;m wide, sometimes inflated at septa. \u003cem\u003eHymenial cystidia\u0026nbsp;\u003c/em\u003eabsent.\u003cem\u003ePileipellis\u003c/em\u003e an ixocutis of thin-walled, 3\u0026ndash;8 \u0026mu;m wide, subparallel to loosely intertwined cylindrical hyphae, sometimes inflated at septa, faintly yellowish; terminal ascending (erect) elements rare to frequent, immersed in a gelled matrix, mostly in clusters, sinuous, subcylindrical with rounded apex, 2\u0026ndash;4 \u0026mu;m wide, often multi-articulated into multiple segments; pigment absent to pale yellow, parietal or rarely minutely incrusting some hyphae. \u003cem\u003eSubpellis\u003c/em\u003eof subparallel to loosely intertwined 3\u0026ndash;10 \u0026mu;m wide hyaline hyphae. \u003cem\u003eThromboplerous hyphae\u003c/em\u003enot observed. \u003cem\u003eStipitipellis\u003c/em\u003econsisting of parallel, faintly yellowish, 2\u0026ndash;4 \u0026mu;m wide cylindrical repent hyphae, occasionally producing clusters of erect, clavate, subglobose to cylindrical (and then multi-articulate) 4\u0026ndash;8 \u0026micro;m wide hyaline caulocystidioid hyphae (elements). \u003cem\u003eStipititrama\u003c/em\u003e of hyaline, short cylindrical, slightly thick-walled (wall up to 0.5 \u0026mu;m thick), 3\u0026ndash;7 \u0026mu;m wide hyphae. \u003cem\u003eClamp connections\u003c/em\u003e absent.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHabitat and distribution\u003c/em\u003e: Boreal forests of \u003cem\u003ePicea abies\u003c/em\u003e, gregarious on needle litter, August-October (European collections), so far known from Finland, Sweden and North America (USA, Oregon).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMaterial examined\u003c/em\u003e: \u003cstrong\u003eFinland\u003c/strong\u003e, Kainuu, Paltamo, Kontiom\u0026auml;ki, mesic \u003cem\u003ePicea abies\u003c/em\u003e forest, 14 September 2008, \u003cem\u003eleg. I. Kyt\u0026ouml;vuori\u003c/em\u003e, H6032677 (H); Suomussalmi, Lohivaara, herb-rich \u003cem\u003ePicea abies\u003c/em\u003e forest on needle litter, 29 August 2019, \u003cem\u003eleg. T. Kekki (3988) \u0026amp; T. Helo\u003c/em\u003e, OULU:GAJ.11332; Koillismaa, Taivalkoski, Katajavaara, old mesic \u003cem\u003ePicea abies\u003c/em\u003e forest with damp grass-herb depressions, on needle litter, 02 September 2008, \u003cem\u003eleg. I. Kyt\u0026ouml;vuori\u003c/em\u003e, H6082249 (H); Per\u0026auml;-Pohjanmaa, Kemij\u0026auml;rvi, Pyh\u0026auml;tunturi, fairly old \u003cem\u003ePicea abies\u003c/em\u003e forest, 27 August 2008, \u003cem\u003eleg.\u003c/em\u003e \u003cem\u003eI. Kyt\u0026ouml;vuori\u003c/em\u003e, H6032673 (H); Rovaniemi, Kylm\u0026auml;oja, mesic \u003cem\u003ePicea abies\u0026nbsp;\u003c/em\u003eforest by a stream, on needle litter, 04 October 2023, \u003cem\u003eleg. T. Kekki (6650)\u003c/em\u003e, OULU:GAJ.17963; Tervola, Raem\u0026auml;ki, calciferous \u003cem\u003ePicea abies\u0026nbsp;\u003c/em\u003eforest, on needle litter, 04 October 2023, \u003cem\u003eleg. T. Kekki (6670)\u003c/em\u003e, OULU:GAJ.17962 (\u003cstrong\u003eHolotype\u003c/strong\u003e, Isotype AMB 20536); ibid. 04 October 2023, \u003cem\u003eleg. T. Kekki (6666)\u003c/em\u003e, OULU:GAJ.17965; ibid. 04 October 2023, \u003cem\u003eleg.\u003c/em\u003e \u003cem\u003eT. Kekki\u003c/em\u003e (6668), OULU:GAJ.17964. \u003cstrong\u003eSweden\u003c/strong\u003e, J\u0026auml;mtland, \u0026Aring;re, mesic \u003cem\u003ePicea abies\u003c/em\u003e forest, 23 August 2006, leg. \u003cem\u003eM. Toivonen \u0026amp; I. Kyt\u0026ouml;vuori\u003c/em\u003e, H7001634 (H).\u003c/p\u003e\n\u003cp\u003eNotes \u0026ndash; The species grows gregarious, in autumn, in the thick needle litter under \u003cem\u003ePicea abies\u003c/em\u003e. It is probably circumpolar as there are collections from Finland, Sweden, and North America. It is rare and prefers old growth \u003cem\u003ePicea\u003c/em\u003e forests. \u003cem\u003eLulesia fallacioides\u003c/em\u003e can be confused with \u003cem\u003eLeucopaxillus alboalutaceus\u003c/em\u003e (F.H. M\u0026oslash;ller \u0026amp; Jul. Sch\u0026auml;ff.) F.H. M\u0026oslash;ller, \u003cem\u003eRipartites\u0026nbsp;\u003c/em\u003eP. Karst.\u003cem\u003e\u0026nbsp;\u003c/em\u003espp. and centrally-stipitate \u003cem\u003eClitopilus\u003c/em\u003e (Fr. ex Rabenh.) P. Kumm. species, but its basidiospore structure is diagnostic. \u003cem\u003eLulesia fallax\u003c/em\u003e is morphologically very similar, but it grows usually in deciduous forests and grasslands, has a xerocutis (versus an ixocutis), and narrower oblong basidiospores (4.1 \u0026mu;m and Q = 1.60 versus 4.7 \u0026mu;m and Q = 1.38, on average) (Vizzini et al. 2023). The Swedish collection named \u003cem\u003eRhodocybe fallax\u003c/em\u003e in von Bonsdorff et al. (2014), due to its habitat, viscid pileus and basidiospore shape is surely attributable to \u003cem\u003eL. fallacioides\u003c/em\u003e. The short description of \u003cem\u003eR\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e included in Funga Nordica by Noordeloos (2012), with basidiospores 5\u0026ndash;7 \u0026times; 3\u0026ndash;4 \u0026micro;m, pileus 10\u0026ndash;40 mm, stipe 2\u0026ndash;7 mm wide, in deciduous woods, scrubs, calcareous sand dunes, in litter under \u003cem\u003eSyringa\u003c/em\u003e, \u003cem\u003eAlnus\u003c/em\u003e, indicates that the true \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e is present in Scandinavia. Our analyses (Figs. 1\u0026ndash;2) show that based on molecularly confirmed collections, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e is present in Nordic areas as Finland (H6032675, Fig. 4a) and Estonia. The Chinese \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eneofallax\u003c/em\u003e (see below) and the Swiss \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e (see below) are mainly distinguished from \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e by smaller basidiomes, a xerocutis as pileipellis, smaller basidiospores, and different habitat (Tab. 2, Liu et al. 2024).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eLulesia blancii\u003c/em\u003e (Maire) Vizzini, Consiglio, P. Alvarado, Angelini \u0026amp; M. Marchetti is a xerophilic species from Mediterranean basin distinguished by a dry alutaceous ochre, chamois, grey-brown pileus, lamellae with evident yellow-ochre tinges, and shorter basidiospores (5.5 \u0026micro;m long on average) (Contu 1999; Eyssartier and Roux 2011; Ivaldi et al. 2023; Vizzini et al. 2023).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia parvifallax\u0026nbsp;\u003c/em\u003e\u003c/strong\u003eRickmann, Vizzini, Consiglio \u0026amp; A. Gross \u003cstrong\u003e\u003cem\u003esp. nov.\u0026nbsp;\u003c/em\u003e\u003c/strong\u003eFigs. 3d\u0026ndash;e, 6\u003c/p\u003e\n\u003cp\u003eMycoBank MBXXXXXXX\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eEtymology\u003c/em\u003e: from the Latin adjectives \u003cem\u003eparvus\u003c/em\u003e (small) and \u003cem\u003efallax\u003c/em\u003e (false, fallacious), due to its similarity to \u003cem\u003eLulesia fallax\u003c/em\u003e, from which it differs especially by having smaller basidiomes and smaller basidiospores.\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDiagnosis\u003c/em\u003e:\u003cem\u003e\u0026nbsp;\u003c/em\u003eIt differs from \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u0026nbsp;\u003c/em\u003eby smaller basidiomes and smaller basidiospores.\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHolotype\u003c/em\u003e (here designated): G00586348\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePileus\u003c/em\u003e 15\u0026ndash;35(\u0026ndash;45) mm diam., convex, soon applanate or depressed at center, getting irregular when old. Margin involute when young, later more or less undulating, often crenulate, not striate. Surface smooth or slightly pruinose, dry, white, sometimes with cream tinges, not hygrophanous, often concentrically cracking and showing the darker context. \u003cem\u003eLamellae\u003c/em\u003e L = 25\u0026ndash;40, l = 1\u0026ndash;3, moderately crowded, decurrent, anastomosing, 1.5\u0026ndash;3 mm broad, whitish, later tinged pinkish cream, edge smooth, concolorous. \u003cem\u003eStipe\u003c/em\u003e 10\u0026ndash;35 \u0026times; 2\u0026ndash;5 mm, central or slightly eccentric attached, cylindrical, usually tapering downwards, smooth, solid, ivory white to pale ochre beige, often with a tinge of pink towards the base, with white rhizomorphs at the base. \u003cem\u003eContext\u003c/em\u003e whitish. Smell not distinctive, taste strongly bitter. No part of the basidiome staining black. KOH on dried basidiome surfaces negative. Spore-print cream pink.\u003c/p\u003e\n\u003cp\u003eBasidiospores (4.7\u0026ndash;)5.0\u0026ndash;\u003cem\u003e5.5\u003c/em\u003e\u0026ndash;5.9(\u0026ndash;6.2) \u0026times; (3.2\u0026ndash;)3.5\u0026ndash;\u003cem\u003e3.9\u003c/em\u003e\u0026ndash;4.3(\u0026ndash;4.6) \u0026micro;m [60/2/1],\u003cbr\u003eQ = (1.26\u0026ndash;)1.34\u0026ndash;\u003cem\u003e1.43\u003c/em\u003e\u0026ndash;1.55(\u0026ndash;1.63), V= (25.7\u0026ndash;)34.3\u0026ndash;\u003cem\u003e43.9\u003c/em\u003e\u0026ndash;57.9(\u0026ndash;66.5), subamygdaliform in side view, ovoid-ellipsoid in frontal view, angular in polar view (5\u0026ndash;7 facets), weakly nodulose-pustulate to verrucose in all views, rounded to slightly attenuated or subconical apically, often adhering in tetrads, usually thin-walled, rarely somewhat thick-walled, hilar appendage evident, up to 1 \u0026micro;m long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cyanophilic, pale yellow in water, yellow in Melzer\u0026rsquo;s. \u003cem\u003eBasidia\u003c/em\u003e 22\u0026ndash;30 \u0026times; 6\u0026ndash;8 \u0026micro;m, cylindrical to clavate-cylindrical, usually with guttulate contents, mainly tetrasporic, rarely bisporic, sterigmata up to 6 \u0026micro;m long. \u003cem\u003eSubhymenium\u003c/em\u003e 30\u0026ndash;50 \u0026micro;m thick, consisting of irregular elements, up to 12 \u0026micro;m wide. \u003cem\u003eHymenophoral trama\u003c/em\u003e regular to subregular of parallel to slightly interwoven, cylindrical or inflated, thin-walled, hyaline, 4\u0026ndash;14 \u0026micro;m wide hyphae. \u003cem\u003eHymenial cystidia\u0026nbsp;\u003c/em\u003eabsent. \u003cem\u003ePileipellis\u003c/em\u003e a xerocutis of thin-walled, 3\u0026ndash;6 \u0026micro;m wide, subparallel to loosely intertwined cylindrical hyphae, sometimes inflated at septa, terminal elements inconspicuous, cylindrical, subclavate or subfusiform, with rounded apex, pigment absent or parietal, not incrusting, pale yellow. \u003cem\u003eSubpellis\u003c/em\u003e poorly differentiated, of subparallel to loosely intertwined 3\u0026ndash;6 \u0026micro;m wide hyaline hyphae. \u003cem\u003eThromboplerous hyphae\u003c/em\u003enot observed. \u003cem\u003eStipitipellis\u003c/em\u003econsisting of parallel, faintly yellowish, 3\u0026ndash;5 \u0026micro;m wide cylindrical repent hyphae, terminal elements hardly differentiated. \u003cem\u003eStipititrama\u003c/em\u003e of hyaline, cylindrical, thin-walled, 5\u0026ndash;9 \u0026micro;m wide hyphae. \u003cem\u003eClamp connections\u003c/em\u003e absent.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHabitat and distribution\u003c/em\u003e: Dry grassland on sandy, calcareous soil, accompanied by \u003cem\u003ePinus sylvestris\u003c/em\u003e and \u003cem\u003eQuercus pubescens\u003c/em\u003e. So far only known from the type locality.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMaterial examined\u003c/em\u003e: \u003cstrong\u003eSwitzerland\u003c/strong\u003e, Canton of Valais, municipality of Leuk, 572 m a.s.l., sunny, dry area on sandy, calcareous soil; the ground covered with mosses and lichens; nearby vegetation includes \u003cem\u003ePinus sylvestris, Quercus pubescens\u003c/em\u003e, and numerous bushes of \u003cem\u003eBerberis vulgaris\u003c/em\u003e; other fungal species observed in the vicinity were \u003cem\u003eCalocybe carnea\u003c/em\u003e (Bull.) Donk\u003cem\u003e, Clitocybe barbularum\u003c/em\u003e (Romagn.) P.D. Orton\u003cem\u003e, Entoloma coracis\u003c/em\u003e Brandrud, Dima, Noordel., G.M. Jansen \u0026amp; Vila\u003cem\u003e, Infundibulicybe glareosa\u003c/em\u003e (R\u0026ouml;llin \u0026amp; Monthoux) Harmaja\u003cem\u003e, Lepiota pseudolilacea\u003c/em\u003e Huijsman\u003cem\u003e, Lichenomphalia ericetorum\u003c/em\u003e (Pers.) Voitk, Thorn \u0026amp; I. Saar\u003cem\u003e\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;Tulostoma melanocyclum\u0026nbsp;\u003c/em\u003eBres.; 19 October 2024, \u003cem\u003eleg. R. Rickmann\u003c/em\u003e (private herbarium no. RR24.326), G00586348 (\u003cstrong\u003eHolotype\u003c/strong\u003e).\u003c/p\u003e\n\u003cp\u003eNotes \u0026ndash; \u003cem\u003eLulesia parvifallax\u003c/em\u003e is so far only known from the type locality in Switzerland. Its precise habitat requirements still need to be clarified through additional collections.\u003c/p\u003e\n\u003cp\u003eIn the field, reliable distinction from closely related species is hardly possible. Apart from the habitat, only the rather small basidiomes with slender stipes may serve as an indication of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e. However, microscopic differentiation is possible; see also Table 2.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eLulesia\u003c/em\u003e \u003cem\u003efallax\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e differ by having slightly larger basidiospores, usually longer than 6 \u0026micro;m (Vizzini et al. 2023 and Tab. 2). Furthermore, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e has an ixocutis. The recently described \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eneofallax\u003c/em\u003e from China can be distinguished by very minute stipes (only 1\u0026ndash;2 mm wide) and a lower spore quotient (Q = 1.2). So far, this species has not been recorded in Europe (Liu et al. 2025).\u003c/p\u003e\n\u003cp\u003eDespite the morphological similarity of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e with the other white-capped species, it is molecularly quite distant from these in the multigene analysis (Fig. 1) while its closest relative is \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eblancii\u003c/em\u003e (with an ITS sequence similarity of approximately 91%). \u003cem\u003eLulesia blancii\u003c/em\u003e shares with \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u0026nbsp;\u003c/em\u003esimilar basidiospore dimensions but it has usually significantly larger basidiomes (pileus up to 70(\u0026ndash;100) mm broad), darker pileus colours, and is so far only known from Mediterranean regions (Ivaldi et al. 2023; Vizzini et al. 2023).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia neofallax\u003c/em\u003e\u003c/strong\u003e(W.H. Lu, Karun. \u0026amp; S. Tibpromma)Vizzini, Consiglio, \u003cstrong\u003e\u003cem\u003ecomb. nov.\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eMycoBank MBXXXXXXX\u003c/p\u003e\n\u003cp\u003eBasionym: \u003cem\u003eClitocella neofallax\u003c/em\u003e W.H. Lu, Karun. \u0026amp; S. Tibpromma, in Liu et al., Mycology 16(1): 26. (2025) [2024] [MB#571612]\u003c/p\u003e\n\u003cp\u003eIn our analysis \u003cem\u003eClitocella neofallax\u003c/em\u003e is phylogenetically part of \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e (Figs. 1\u0026ndash;2) and it is combined above in \u003cem\u003eLulesia\u003c/em\u003e accordingly. The species is distinguished among the others within the \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e complex by gracile basidiomes, pileus 10\u0026ndash;30 mm wide, dry, low convex, sometimes infundibuliform, with a shallow depression at the centre, stipe 10‒25 \u0026times; 1‒2 mm, lamellae with evident yellowish hues even when young, basidiospores (4.0\u0026ndash;)4.8\u0026ndash;6.3 \u0026times; (3.5\u0026ndash;)4\u0026ndash;5 \u0026mu;m, Q= 1.2 and very short basidia, 15‒23.5 \u0026times; 6‒9 \u0026mu;m (Liu et al. 2025). It is so far known only from China (Yunnan Province, Qujing City, Qujing Normal University) on soil associated with bamboo roots.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003esubgen.\u003cem\u003e\u0026nbsp;Lulesia\u0026nbsp;\u003c/em\u003eSinger, Fl. Neotrop., Monogr. 3: 16 (1970)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAutonym\u003c/p\u003e\n\u003cp\u003eType: \u003cem\u003eArmillariella densifolia\u003c/em\u003e Singer, in Singer \u0026amp; Digilio, Lilloa 25: 72 (1952) [1951]\u003c/p\u003e\n\u003cp\u003ePileus usually pale grey, grey, dark grey, brown, violaceous black, basidiome surfaces unchanging or turning grey or black when bruised or with age, usually with a positive, reddish, KOH reaction, hymenophoral trama usually irregular (intertwined hyphae), and basidiospores globose, subglobose, broadly ellipsoid to ellipsoid, which are obscurely to clearly angular, and weakly to clearly pustulate.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia alachuana\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e(Murrill) Singer, Fl. Neotrop., Monogr. 3: 17 (1970) Figs. 3i\u0026ndash;j, 7\u0026ndash;8\u003c/p\u003e\n\u003cp\u003eBasionym: \u003cem\u003eClitocybe alachuana\u003c/em\u003e Murrill, Proc. Fla Acad. Sci. 7(2/3): 107 (1945) [1944]\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHolotype\u003c/em\u003e: FLAS F-17903, USA, Florida, Alachua, Prairie Creek Hammock, on dead leaves, 15 July 1938, \u003cem\u003eleg. West, Arnold, and Murrill\u003c/em\u003e, \u003cem\u003edet. Murrill\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eEpitype\u003c/em\u003e: FLAS F-61088 (Designated by Vizzini, Alvarado, Consiglio, Angelini \u0026amp; Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 5. 2024). Registration Identifier 10017392. MycoBank MBT10017392\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ePileus\u003c/em\u003e 20\u0026ndash;80 mm diam., at first convex, soon depressed; surface hygrophanous, dry, subglabrous, at first covered with a minute white soon disappearing pruina (which often fragments concentrically) that persists on the margin, grey avellaneous, tan (5C5\u0026ndash;7), brown (4E6\u0026ndash;7), dark brown (1F5\u0026ndash;7, 3F4\u0026ndash;8, 4F6\u0026ndash;7), sometimes with purplish-grey tones (7F2\u0026ndash;5), margin incurved, uneven, undulate, lobed. \u003cem\u003eLamellae\u003c/em\u003e long decurrent, narrow, very close (crowded) L= 40\u0026ndash;70, l= 1\u0026ndash;3, some forked halfway, at the base and at the apex, entire, at first white, pale yellowish (3A7\u0026ndash;8) when old, with pinkish tones, never turning grey black on handling or bruising, with an even, sinuous, concolorous or paler edge. \u003cem\u003eStipe\u003c/em\u003e 15\u0026ndash;30 \u0026times; 5\u0026ndash;8 (\u0026ndash;10) mm, usually equal, solitary, often flared at the apex, rarely attenuated at the base, at first white, then beige (5A3), cream avellaneous (6A3\u0026ndash;4), white tomentose, especially at the base, with white rhizomorphs at the base. \u003cem\u003eContext\u003c/em\u003e very thin, white, unchanging; \u003cem\u003esmell\u003c/em\u003e not recorded (of light anise in FLAS-F-71949-iNaturalist-183000711), \u003cem\u003etaste\u003c/em\u003e very bitter. Chemical spot-test reactions: KOH on pileus surface negative.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eBasidiospores\u003c/em\u003e (4.3\u0026ndash;)4.8\u0026ndash;\u003cem\u003e5.1\u003c/em\u003e\u0026ndash;5.4(\u0026ndash;6.0) \u0026times; (3.6\u0026ndash;)4.2\u0026ndash;\u003cem\u003e4.5\u003c/em\u003e\u0026ndash;4.7(\u0026ndash;5.2) \u0026micro;m [192/4/4], Q = (1.00\u0026ndash;)1.06\u0026ndash;\u003cem\u003e1.14\u003c/em\u003e\u0026ndash;1.21(\u0026ndash;1.36), V = (33.3\u0026ndash;)45.8\u0026ndash;\u003cem\u003e54.0\u003c/em\u003e\u0026ndash;62.3(\u0026ndash;75.6) \u0026micro;m\u003csup\u003e3\u003c/sup\u003e, globose, subglobose up to broadly ellipsoid in side and frontal view, clearly angular (9\u0026ndash;12 facets) especially when young, obscurely angular when mature, with small pustules or bumps, hilar appendage up to 1 \u0026mu;m long, contents smooth, uni- to pluriguttulate (guttulae greenish in water), wall cianophilic, faintly congophilic, hyaline to pale yellow in water, yellow in Melzer\u0026rsquo;s (inamyloid). \u003cem\u003eBasidia\u003c/em\u003e 18\u0026ndash;30(\u0026ndash;35) \u0026times; 6.5\u0026ndash;8 \u0026mu;m, cylindrical-clavate, tetrasporic, rarely bisporic, sterigmata up to 5 \u0026mu;m long, usually with guttulate greenish contents. \u003cem\u003eSubhymenium\u003c/em\u003e 20\u0026ndash;40 \u0026mu;m thick, consisting of textura intricata type short elements, 2\u0026ndash;5 \u0026mu;m wide. \u003cem\u003eHymenophoral trama\u003c/em\u003e subregular, hyphae subparallel to intertwined, cylindrical, 2\u0026ndash;8 \u0026mu;m wide, thick-walled (wall up to 0.6 \u0026micro;m thick), hyaline to faintly cream. \u003cem\u003eHymenial cystidia\u003c/em\u003e usually absent. In the collection FLAS-F-61088, cystidioid hyaline and thin-walled elements, subcylindrical, sinuous, 20\u0026ndash;50 \u0026times; 2\u0026ndash;4 \u0026micro;m, are occasionally present on the lamellar edge (Fig. 7f). \u003cem\u003ePileipellis\u003c/em\u003e as a xerocutis with cylindrical, mostly intertwined hyphae, 2.5\u0026ndash;6(\u0026ndash;7) \u0026mu;m wide, thick-walled (wall up to 0.6 \u0026micro;m thick), hyaline, smooth, the more superficial ones reclined to ascendant, hyaline or with a pale yellow cytoplasmatic pigment, pileocystidia not observed; \u003cem\u003ethromboplerous hyphae\u003c/em\u003e absent. \u003cem\u003eSubpellis\u003c/em\u003e of subparallel to loosely intertwined 3\u0026ndash;8 \u0026mu;m wide hyaline hyphae. \u003cem\u003eStipitipellis\u003c/em\u003e as a cutis consisting of subparallel to intertwined loose hyphae, 2\u0026ndash;5 \u0026micro;m wide, faintly yellowish; \u003cem\u003ecaulocystidia\u003c/em\u003e as dense tufts of reclining to erect hyphoid elements, often multiseptate, straight to wavy, short to long, 30\u0026ndash;100(\u0026ndash;150) \u0026times; 2\u0026ndash;4 \u0026micro;m, with an obtuse to acute apex, sometimes also subcapitate, smooth or with an epiparietal pigment in patches. \u003cem\u003eStipititrama\u003c/em\u003e made up of parallel subcylindrical, 3\u0026ndash;7 \u0026micro;m wide, hyaline hyphae, thin- to thick-walled (wall up to 0.5 \u0026micro;m thick).\u003cem\u003e\u0026nbsp;Clamp connections\u003c/em\u003e absent.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMaterial examined\u003c/em\u003e: \u003cstrong\u003eUSA\u003c/strong\u003e, Florida, Alachua, Prairie Creek Hammock, 29.598463 -82.247989 +-1600 m, on dead leaves, 15 July 1938, \u003cem\u003eleg. West\u003c/em\u003e, \u003cem\u003eArnold\u003c/em\u003e \u0026amp; \u003cem\u003eMurrill\u003c/em\u003e, FLAS-F-17903 (\u003cstrong\u003eHolotype\u003c/strong\u003e, as \u003cem\u003eClitocybe\u003c/em\u003e \u003cem\u003ealachuana\u003c/em\u003e); Florida, Alachua, Gainesville, 29.66636 -82.32993 +-8000 m, trash in rich hammock, 08 August 1939, \u003cem\u003eleg.\u003c/em\u003e \u003cem\u003eE\u003c/em\u003e. \u003cem\u003eWest\u003c/em\u003e, \u003cem\u003edet.\u003c/em\u003e \u003cem\u003eWA Murrill\u003c/em\u003e, FLAS-F-19879 (as \u003cem\u003eC. alachuana\u003c/em\u003e); Florida, Putnam County, Ordway-Swisher Biological Station, Mill Creek Swamp Bridge, 30.80138 -86.292863, on decorticate wood, \u003cem\u003eQuercus\u003c/em\u003e and palm tree dominated swamp, 06 July 2017, \u003cem\u003eleg\u003c/em\u003e. \u003cem\u003eD Borland \u0026amp; B Kaminsky\u003c/em\u003e, \u003cem\u003edet. ME Smith\u003c/em\u003e (as cf. \u003cem\u003eClitocella\u003c/em\u003e), FLAS-F-61088 (\u003cstrong\u003eEpitype\u003c/strong\u003e), duplo in CORT014753, \u003cem\u003edet. TJ Baroni\u003c/em\u003e (as \u003cem\u003eLulesia alachuana\u003c/em\u003e); Florida, Melrose, Putnam County, Ordway-Swisher Biological Station, 29.71900833 -81.97166333, on the ground in highly decayed litter and also very decayed wood of hardwoods, 12 September 2023, \u003cem\u003eleg. F Sheffer\u003c/em\u003e, FLAS-F-71949-iNaturalist-183000711(https://www.inaturalist.org/observations/183000711) (as \u003cem\u003eClitocella\u003c/em\u003e sp.).\u003c/p\u003e\n\u003cp\u003eNotes \u0026ndash; \u003cem\u003eLulesia alachuana\u003c/em\u003e, originally described from Florida (Murrill 1944, Alachua County, \u003cem\u003einde nomen\u003c/em\u003e, from which the species name derives), is so far known only from Florida (Tab. 1 and Fig. 1). The present full description of the species is the first to be provided after the original one by Murrill (1944). Due to its greyish brown pileus, pileipellis as a cutis, subglobose, slightly angular, undulate pustulate or nearly smooth spores, a hymenophoral trama of interwoven hyphae, and bitter taste (Murrill 1944; Bigelow 1982; our observations), \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ealachuana\u003c/em\u003e is also morphologically a good member of \u003cem\u003eLulesia\u0026nbsp;\u003c/em\u003esubgen. \u003cem\u003eLulesia\u003c/em\u003e, as molecularly supported in the multigene analysis (Fig. 1) where it is sister to a strongly supported clade (BP = 95%, PP = 1.00) consisting of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eobscura\u0026nbsp;\u003c/em\u003eand \u003cem\u003eL\u003c/em\u003e. \u003cem\u003epopinalis\u0026nbsp;\u003c/em\u003e(all with greyish colours, irregular hymenophoral trama and red reaction to KOH). Because of its subglobose spores 5.1 \u0026times; 4.5 \u0026mu;m, Q = 1.14 on average, and growth on forest (hardwood) litter \u003cem\u003eL. alachuana\u0026nbsp;\u003c/em\u003eseems quite close to \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u0026nbsp;\u003c/em\u003eand in fact it was considered a later synonym of the latter by some authors [e.g., T. Baroni\u0026rsquo;s handwritten notes (1978) accompanying the holotype collection of\u003cem\u003e\u0026nbsp;Clitocybe\u003c/em\u003e \u003cem\u003ealachuana\u003c/em\u003e (FLAS-F-17903), https://www.mycoportal.org/portal/collections/individual/index.php?occid=604249\u0026amp;clid=0; Baroni (1981); Bigelow (1982, 1985); Singer (1986)]. It morphologically seems to differ from \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e mainly by a darker and hygrophanous pileus (brown, dark brown versus white, pale cream, beige-ochraceous), a non-blackening context, a pileus surface with negative KOH reaction (greyish brown versus red) and different nrDNA sequences (Fig. 1; Vizzini et al. 2024 and our observations).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia colorata\u003c/em\u003e\u003c/strong\u003e(L. Fan \u0026amp; N. Mao) T.J. Baroni, N. Niveiro \u0026amp; B.E. Lechner, in Baroni, Lechner \u0026amp; Niveiro, Index Fungorum 566: 1 (2023) Figs. 3g\u0026ndash;h, 4b\u003c/p\u003e\n\u003cp\u003eBasionym: \u003cem\u003eClitocella colorata\u003c/em\u003e L. Fan \u0026amp; N. Mao, in Mao, Lv, Xu, Zhao \u0026amp; Fan, MycoKeys 88: 161 (2022)\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHolotype\u003c/em\u003e: BJTC FM1891\u003c/p\u003e\n\u003cp\u003eFor full descriptions (as \u003cem\u003eClitocella\u003c/em\u003e \u003cem\u003ecolorata\u003c/em\u003e) see Mao et al. (2022) and Vizzini et al. (2023)\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMaterial examined\u003c/em\u003e: \u003cstrong\u003eFinland\u003c/strong\u003e, Etel\u0026auml;-Karjala, Kouvola, somewhat springfed spruce-hardwood swamp, 19 September 1994, \u003cem\u003eleg. I. Kyt\u0026ouml;vuori\u003c/em\u003e (as \u003cem\u003eRhodocybe\u003c/em\u003e cf. \u003cem\u003emundula\u003c/em\u003e), \u003cem\u003edet. T. Kekki\u003c/em\u003e, H6042258 (H). \u003cstrong\u003eItaly\u003c/strong\u003e. Lombardia, Pavia, Brallo di Pregola, under \u003cem\u003eFagus sylvatica\u003c/em\u003e, 15 October 2022, \u003cem\u003eleg. M. Carbone\u003c/em\u003e, TUR-A 216670; Emilia-Romagna, Piacenza, Ferriere, Loc. Le Moline, mixed forest with \u003cem\u003eQuercus cerris\u003c/em\u003e, \u003cem\u003eQ\u003c/em\u003e. \u003cem\u003epubescens\u003c/em\u003e and \u003cem\u003eOstrya carpinifolia\u003c/em\u003e, 12 October 2019, \u003cem\u003eleg. et det. M. Carbone \u0026amp; F. Calledda\u003c/em\u003e (as \u003cem\u003eClitocella\u003c/em\u003e \u003cem\u003emundula\u003c/em\u003e), TUR-A 216671.\u003c/p\u003e\n\u003cp\u003eNotes \u0026ndash; \u003cem\u003eLulesia colorata\u003c/em\u003e (as \u003cem\u003eClitocella\u003c/em\u003e \u003cem\u003ecolorata\u003c/em\u003e) has recently been described from China (Shanxi province, North China or Huabei) on soil or rotten wood in coniferous (\u003cem\u003ePinus\u003c/em\u003e) or broad-leaved (\u003cem\u003eQuercus\u003c/em\u003e) forest (Mao et al. 2022). The Chinese authors highlighted that also a northamerican collection named as \u003cem\u003eC\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e (TJB7599 (AFTOL-ID 521) USA, NY) clustered within the \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e clade. In the original description, no reference is made to a possible colour change of the basidiome surfaces, its smell and taste are not reported, and the hymenophoral trama is indicated as regular (Mao et al. 2022). In their monographic work on the genus \u003cem\u003eClitocella\u003c/em\u003e in Europe, Vizzini et al. (2023), based on molecular data and new collections (from Italy-Veneto and Estonia in mixed coniferous forests) highlighted that \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e has: 1) a wider geographical distribution, and it is also present in India, South Korea, USA (Arkansas, Indiana, New York, Tennessee, Wisconsin) and in Europe (France, Italy-Veneto, and Estonia) (Fig. 1); 2) pileus, lamellae, and stipe surface stain strongly black when bruised or in age, smell is farinaceous, taste is bitter, and the hymenophoral trama is subregular at first but then irregular and composed of interwoven hyphae. Our present analysis allow to extend the geographical distribution of this species to Finland and the Italian regions of Lombardy and Emilia-Romagna and to confirm the additional features provided by Vizzini et al. (2023), including also the blackening of basidioma surfaces. Consequently, an emended combination of characters circumscribes \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata\u0026nbsp;\u003c/em\u003esuch as clitocyboid basidiomes (strongly depressed pileus), a usually pale-coloured pileus surface (white to yellowish white, greyish white to greyish brown, pink-white), the surfaces of the basidiome turning black when old or on handling and showing no reddish reaction in KOH, farinaceous smell and bitter taste, (4.7\u0026ndash;)5.1\u0026ndash;\u003cem\u003e5.5\u003c/em\u003e\u0026ndash;5.9(\u0026ndash;6.8) \u0026times; (3.7\u0026ndash;)4.1\u0026ndash;\u003cem\u003e4.3\u003c/em\u003e\u0026ndash;4.6(\u0026ndash;5.0) \u0026micro;m, (globose) subglobose to broadly ellipsoid basidiospores slightly angular and with minute pustules or bumps, hyphae of pileipellis with pale yellow to yellowish brown intracellular and/or parietal pigment, and the growth in coniferous or angiospermous forests (Mao et al. 2022; Vizzini et al. 2023; our observations).\u003c/p\u003e\n\u003cp\u003eMorphologically, \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e can be easily confused with \u003cem\u003eC\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e and \u003cem\u003eC\u003c/em\u003e. \u003cem\u003epopinalis\u003c/em\u003e which, however, have dried basidiome surfaces producing a reddish reaction with KOH (Baroni 1981; Moreau 1997; Kluting et al. 2014; Vizzini et al. 2023). Indeed, many collections that clustered in the \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e clade (Fig. 1) were originally identified as \u003cem\u003eC\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e or \u003cem\u003eC\u003c/em\u003e. \u003cem\u003epopinalis\u003c/em\u003e (Fig. 1 and Tab. 1). Additionally, \u003cem\u003eC. mundula\u003c/em\u003e differs in the obscurely angular basidiospores with indistinct pustules or bumps (Baroni 1981; Neukom 1994; Moreau 1997) and \u003cem\u003eC. popinalis\u003c/em\u003e in basidiome surfaces usually unchanging on handling, a dark-coloured pileus slightly depressed only in senescing basidiomes, its association with grasses, and slightly broader and longer basidiospores (Baroni 1981; Moreau 1997; Overall 2011; Kluting et al. 2014; Jian et al. 2020a).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLulesia solaris\u003c/em\u003e\u003c/strong\u003e(Musumeci, Consiglio \u0026amp; Vizzini) Musumeci, Consiglio \u0026amp; Vizzini, in Vizzini, Alvarado, Consiglio, Angelini \u0026amp; Marchetti, Boll. Assoc. Micol. Ecol. Romana 39(3): 8. (2024) [2023]. Fig. 4f\u003c/p\u003e\n\u003cp\u003eBasionym: \u003cem\u003eClitocella solaris\u003c/em\u003e Musumeci, Consiglio \u0026amp; Vizzini, in Vizzini et al., Persoonia 50: 144 (2023)\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eHolotype\u003c/em\u003e: LUG 19882\u003c/p\u003e\n\u003cp\u003eFor a full description (as \u003cem\u003eC\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e) see Vizzini et al. (2023).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMaterial examined\u003c/em\u003e: \u003cstrong\u003eFinland\u003c/strong\u003e, Kittil\u0026auml;n Lappi, Kolari, \u003cem\u003ePinus sylvestris\u003c/em\u003e forest near limestone quarry, 16 August 2001, leg. I. Kyt\u0026ouml;vuori (as \u003cem\u003eRhodocybe\u003c/em\u003e \u003cem\u003ecaelata\u003c/em\u003e), det. T. Kekki (H6043012).\u003c/p\u003e\n\u003cp\u003eNotes \u0026ndash; The species was recently described from France (D\u0026eacute;partement Haut-Rhin, Alsace) as gregarious on moss pads (\u003cem\u003eRhacomitrium canescens\u003c/em\u003e) in open and sunny areas (with seedlings of \u003cem\u003eBetula pubescens\u003c/em\u003e and at some distance (at least 30 m) two \u003cem\u003ePinus sylvestris\u0026nbsp;\u003c/em\u003etrees, on sandy-pebbly, alluvial soil, silty substrate rich in woody debris and carbonate). Until the present work it was so far known only from the \u003cem\u003elocus typicus\u003c/em\u003e (Vizzini et al. 2023). \u003cem\u003eLulesia solaris\u003c/em\u003e is distinguished by a unique combination of characters such as the small-sized basidiomes (pileus not exceeding 10 mm diam), a whitish cream pileus surface with a positive (red) KOH reaction, distant (spaced) lamellae, an unchanging, not staining black context, subfarinaceous smell and taste, absence of hymenial cystidia and clearly angular, subglobose, broadly ellipsoid to ellipsoid basidiospores with evident bumps in Cotton blue. In the present work,a Finnish twenty-four years old collection misidentified as \u003cem\u003eRhodocybe caelata\u0026nbsp;\u003c/em\u003e(H6043012, see above) turned out to be conspecific with \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e (Fig. 1). It was found on calcareous soil and close to \u003cem\u003ePinus sylvestris\u003c/em\u003e trees as the holotype collection. Molecularly, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e is sister (BP = 99 %, PP = 1.00) to a clade formed by \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e and \u003cem\u003eC\u003c/em\u003e. \u003cem\u003eorientalis\u003c/em\u003e (Fig. 1), species which show larger basidiomes, a negative KOH reaction, crowded lamellae, and differently shaped basidiospores (Jian et al. 2020a; Mao et al. 2022; Vizzini et al. 2023 and our observations). \u003cem\u003eClitocella\u003c/em\u003e \u003cem\u003epopinalis\u003c/em\u003e has a larger pileus and stipe, a usually dark-coloured pileus surface (purplish grey, brownish grey, blackish grey) crowded lamellae, and subglobose to broadly ellipsoid wider basidiospores (5.1 \u0026micro;m on average) (Vizzini et al. 2023).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe morphological delimited \u003cem\u003eL. fallax\u003c/em\u003e species complex is part of \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e (pallid \u003cem\u003eClitopilus\u003c/em\u003e-like coloured basidiomes, regular to subregular hymenophoral trama made up of parallel to slightly intertwined cylindrical hyphae, negative KOH reaction, and often adhering in tetrads spores; Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The \u003cem\u003eL. fallax\u003c/em\u003e complex consists of species sharing a bitter taste, a white-pruinose pileus reminding \u003cem\u003eClitocybe\u003c/em\u003e sect. \u003cem\u003eCandicantes\u003c/em\u003e (Qu\u0026eacute;l.) Singer \u0026amp; Digilio species and often in association with coniferous trees. This species complex encompasses \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eneofallax\u003c/em\u003e and the two new species \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e, which are all very similar taxa morphologically differentiated almost only based on the size of the basidiomes and the basidiospores (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These species together with those gravitating around \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e of \u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e, viz. \u003cem\u003eL\u003c/em\u003e. \u003cem\u003epopinalis\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e, could be considered as cryptic taxa, namely species exhibiting shallow morphological differences (morphological stasis, deceleration of morphological evolution), but considerable genetic disparity (Bickford et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Grebenc et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Korhonen et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Struck et al. \u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Korshunova et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Peintner et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Cerca et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Struck and Cerca \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Ekanayaka et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Cryptic species could be considered as the opposite of adaptive radiation (pronounced morphological differences, shallow genetic divergence) (Cerca et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Struck and Cerca \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and, a priori, they could result from recent speciation, parallelism, convergence or stasis (Struck and Cerca \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR131\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Molecular analyses (Vizzini et al. \u003cspan citationid=\"CR145\" class=\"CitationRef\"\u003e2023\u003c/span\u003e, \u003cspan citationid=\"CR147\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e in the present paper) would indicate that the phenomena of evolutionary convergence and parallelism should be excluded for the appearance of cryptospecies in \u003cem\u003eLulesia\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eExamples of morphological stasis are commonplace in other rhodocyboid fungi, viz. the \u003cem\u003eRhodophana nitellina\u003c/em\u003e complex (Dima et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, Buyck et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Papetti \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and \u003cem\u003eRhodocybe gemina\u003c/em\u003e complex (Crous et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Sesli and Vizzini \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Vizzini et al. \u003cspan citationid=\"CR144\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Silva-Filho et al. \u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Dutta et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Sun and Bau \u003cspan citationid=\"CR133\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eAs all the species covered in the present paper show difficulty in morphological distinction, these may have been misdetermined in the past, and a greater scrutiny and increased sequencing of historical and modern collections are now imperative before any conclusions can be drawn regarding more precise distribution and rarity data for each of these species in the world.\u003c/p\u003e\u003cp\u003eThe occurrence of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e in North America (Baroni \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1981\u003c/span\u003e) and The Netherlands (Noordeloos \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e1988\u003c/span\u003e) is based on presumably heterogeneous collections and thus giving rise to collective descriptions (basidiospores 6.5\u0026ndash;8 \u0026times; 4\u0026ndash;5(\u0026ndash;6.5) \u0026micro;m and 5.0\u0026ndash;8.5 \u0026times; 3.5\u0026ndash;5.0 \u0026micro;m-Q 1.4\u0026ndash;2.0, respectively). So, the presence of the true \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e in North America and The Netherlands is still questionable. No sequences in the public databases obtained by Dutch fungal collections clustered within the \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e clade and the \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e complex (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The only sequences from an American collection named \u003cem\u003eC\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e (OKM:25668, USA-Oregon), represent the new species \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallacioides\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Two ITS environmental sequences from USA (Utah) (UDB03879566 and UDB03879567) and two basidiomatal sequences (USA-California and China-Xizang) (PP594830 and PV124053) clustered in a clade (\u003cem\u003eLulesia\u003c/em\u003e sp. 2) within \u003cem\u003eLulesia\u003c/em\u003e subg. \u003cem\u003eParaclitopilus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) which probably represents a distinct not yet described species.\u003c/p\u003e\u003cp\u003eSwiss specimens growing on bare soil of a riparian forest named \u003cem\u003eRhodocybe fallax\u003c/em\u003e (Breitenbach \u0026amp; Kr\u0026auml;nzlin \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1995\u003c/span\u003e), characterized by gracile, diminutive omphaloid basidiomes (pileus 10\u0026ndash;20 mm wide, infundibuliform, stipe 10\u0026ndash;30 \u0026times; 1.5\u0026ndash;3 mm) with basidiospores 7\u0026ndash;8.6 \u0026times; 3.8\u0026ndash;4.8 (Q\u0026thinsp;=\u0026thinsp;1.7\u0026ndash;1.9, V\u0026thinsp;=\u0026thinsp;76) and resembling \u003cem\u003eClitopilus\u003c/em\u003e sect. \u003cem\u003eScyphoides\u003c/em\u003e Singer (Singer \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Jian et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e), are macromorphologically similar to those cited in Baroni (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1981\u003c/span\u003e). They will have to be further studied and molecularly checked to ascertain their taxonomic status.\u003c/p\u003e\u003cp\u003eFinally, the presence of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eobscura\u003c/em\u003e in Finland is here molecularly confirmed for the first time (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and see Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e4\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003eMatteo Carbone (Italy), Balint Dima (Hungary), Leanne P. Sheffer, Matthew E. Smith and Caroline B. Willis (USA, Florida), Andrin Gross (Switzerland), Diana Weckman (Finland) are acknowledged for dried fungal material and sequences and photos of some collections. We thank Balint Dima (Hungary), Kare Liimatainen, Tuula Niskanen and Matti Kulju (Finland) for the sequences produced in the FinBOL project and included in this study.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contribution\u003c/strong\u003e AV and GC conceived the study, IK, TK and RR conducted sampling and provided materials, MM and RR performed all microscopy analyses mentioned in this study, AV performed the laboratory experiments and prepared all figures and tables, and GC analysed the sequence data. AV wrote the manuscript with contributions from all authors. All authors agreed with the submission of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis study has been partly supported by the Finnish Barcode of Life (FinBOL) project which operates under the Finnish Biodiversity Information Facility (FinBIF) Research Infrastructure funded by the Research Council of Finland, and by a grant YM23/5512/2013 from the Finnish Ministry of Environment. The sequencing of \u003cem\u003eLulesia parvifallax\u003c/em\u003e was kindly funded by the Swiss Federal Office for the Environment.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e Sequence data have been deposited in GenBank as given in Table 1 and alignments in Figshare (https://doi.org/10.6084/m9.figshare.29654816).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eEthics approval and consent to participate Not applicable.\u003c/p\u003e\n\u003cp\u003eConsent for publication Not applicable.\u003c/p\u003e\n\u003cp\u003eCompeting interests None\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAgerer R (2018) Subphylum \u003cem\u003eAgaricomycotina\u0026nbsp;\u003c/em\u003eDoweld. In Begerow D, Agerer R, McTaggart A, \u003cem\u003eBasidiomycota\u0026nbsp;\u003c/em\u003eand \u003cem\u003eEntorhizomycota\u003c/em\u003e. A. Engler\u0026apos;s Syllabus of Plant Families, part 1/3 (Frey W, ed). Borntraeger, Stuttgart, pp 130\u0026ndash;444\u003c/li\u003e\n \u003cli\u003eAltschul SF, Gish W, Miller W, Myers EW, Lipman DJ\u003cem\u003e\u0026nbsp;\u003c/em\u003e(1990) Basic local alignment search tool. J Mol Biol\u003cem\u003e\u0026nbsp;\u003c/em\u003e215(3):403\u0026ndash;410\u003c/li\u003e\n \u003cli\u003eAlvarado P, Manj\u0026oacute;n JL, Matheny PB, Esteve-Ravent\u0026oacute;s F (2010) \u003cem\u003eTubariomyces\u003c/em\u003e, a new genus of \u003cem\u003eInocybaceae\u003c/em\u003e from the Mediterranean region. Mycologia 102(6):1389\u0026ndash;1397 https://doi.org/10.3852/10-041\u003c/li\u003e\n \u003cli\u003eAlvarado P, Moreno G, Manj\u0026oacute;n JL (2012) Comparison between \u003cem\u003eTuber gennadii\u003c/em\u003e and \u003cem\u003eT\u003c/em\u003e. \u003cem\u003eoligospermum\u003c/em\u003e lineages reveals the existence of the new species \u003cem\u003eT\u003c/em\u003e. \u003cem\u003ecistophilum\u003c/em\u003e (\u003cem\u003eTuberaceae\u003c/em\u003e, \u003cem\u003ePezizales\u003c/em\u003e). Mycologia 104(4):894\u0026ndash;910 https://doi.org/10.3852/11-254\u003c/li\u003e\n \u003cli\u003eAngelini C, Contu M (2012) \u003cem\u003eLulesia densifolia\u0026nbsp;\u003c/em\u003e(\u003cem\u003eBasidiomycota, Agaricomycetes\u003c/em\u003e) rinvenuta nella Repubblica Dominicana (Caraibi). 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FEMS Microbiol Lett\u003cem\u003e\u0026nbsp;\u003c/em\u003e297(1):24\u0026ndash;30\u003c/li\u003e\n \u003cli\u003eHe M-Q, Zhao R-L, Hyde KD, Begerow D, Kemler M, Yurkov A, McKenzie EHC, Rasp\u0026eacute; O, Kakishima M, S\u0026aacute;nchez-Ram\u0026iacute;rez S, Vellinga EC, Halling R, Papp V, Zmitrovich IV, Buyck B, Ertz D, Wijayawardene NN, Cui B-K, Schoutteten N, Liu X-Z, Li T-H, Yao Y-J, Zhu X-Y, Liu A-Q, Li G-J, Zhang M-Z, Ling Z-L, Cao B, Anton\u0026iacute;n V, Boekhout T, da Silva BDB, De Crop E, Decock C, Dima B, Dutta AK, Fell JW, Geml J, Ghobad-Nejhad M, Giachini AJ, Gibertoni TB, Gorj\u0026oacute;n SP, Haelewaters D, He S-H, Hodkinson BP, Horak E, Hoshino T, Justo A, Lim YW, Menolli N, Me\u0026scaron;ić A, Moncalvo J-M, Mueller GM, Nagy LG, Nilsson RH, Noordeloos M, Nuytinck J, Orihara T, Ratchadawan C, Rajchenberg M, SilvaFilho AGS, Sulzbacher MA, Tkalčec Z, Valenzuela R, Verbeken A, Vizzini A, Wartchow F, Wei T-Z, Wei\u0026szlig; M, Zhao C-L, Kirk PM (2019) Notes, outline and divergence times of \u003cem\u003eBasidiomycota\u003c/em\u003e. Fungal Divers\u003cem\u003e\u0026nbsp;\u003c/em\u003e99(1):105\u0026ndash;367 https://doi.org/10.1007/s13225-019-00435-4\u003c/li\u003e\n \u003cli\u003eHe M-Q, Cao B, Liu F, Boekhout T, Denchev TT, Schoutteten N, Denchev CM, Kemler M, Gorj\u0026oacute;n SP, Begerow D, Valenzuela R, Davoodian N, Niskanen T, Vizzini A, Redhead SA, Ram\u0026iacute;rez-Cruz V, Papp V, Dudka VA, Dutta AK, Garc\u0026iacute;a-Sandoval R, Liu X-Z, Kijpornyongpan T, Savchenko A, Tedersoo L, Theelen B, Trierveiler-Pereira L, Wu F, Zamora JC, Zeng X-Y, Zhou L-W, Liu S-L, Ghobad-Nejhad M, Giachini AJ, Li G-J, Kakishima M, Olariaga I, Haelewaters D, Sulistyo B, Sugiyama J, Svantesson S, Yurkov A, Alvarado P, Anton\u0026iacute;n V, da Silva AF, Druzhinina I, Gibertoni TB, Guzm\u0026aacute;n-D\u0026aacute;valos L, Justo A, Karunarathna SC, Galappaththi MCA, Toome-Heller M, Hosoya T, Liimatainen K, M\u0026aacute;rquez R, Me\u0026scaron;ić A, Moncalvo J-M, Nagy LG, Varga T, Orihara T, Raymundo T, Salcedo I, Silva-Filho AGS, Tkalčec Z, Wartchow F, Zhao C-L, Bau T, Cabarroi-Hern\u0026aacute;ndez M, Cort\u0026eacute;s-P\u0026eacute;rez A, Decock C, De Lange R, Weiss M, Menolli N, Nilsson RH, Fan Y-G, Verbeken A, Gafforov Y, Meiras-Ottoni A, Mendes-Alvarenga RL, Zeng N-K, Wu Q, Hyde KD, Kirk PM, Zhao R-L (2024) Phylogenomics, divergence times and notes of orders in \u003cem\u003eBasidiomycota\u003c/em\u003e. Fungal Divers 126(1):127\u0026ndash;406 https://doi.org/10.1007/s13225-024-00535-w\u003c/li\u003e\n \u003cli\u003eHe X-L, Li T-H, Xi P-G, Jiang Z-D, Shen Y-H (2013) Phylogeny of \u003cem\u003eEntoloma\u0026nbsp;\u003c/em\u003es.l. subgenus \u003cem\u003ePouzarella\u003c/em\u003e, with descriptions of five new species from China. Fungal Divers\u003cem\u003e\u0026nbsp;\u003c/em\u003e58(1):227\u0026ndash;243 https://doi.org/10.1007/s13225-012-0212-7\u003c/li\u003e\n \u003cli\u003eHe Z-M, Yang ZL (2022) The genera \u003cem\u003eBonomyces\u003c/em\u003e, \u003cem\u003eHarmajaea\u0026nbsp;\u003c/em\u003eand \u003cem\u003eNotholepista\u0026nbsp;\u003c/em\u003efrom Northwestern China: two new species and a new record. Mycol Prog\u003cem\u003e\u0026nbsp;\u003c/em\u003e21(2):26 https://doi.org/10.1007/s11557-022-01786-0\u003c/li\u003e\n \u003cli\u003eHofstetter V, Cl\u0026eacute;men\u0026ccedil;on H, Vilgalys R, Moncalvo J-M (2002) Phylogenetic analyses of the \u003cem\u003eLyophylleae\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAgaricales, Basidiomycota\u003c/em\u003e) based on nuclear and mitochondrial rDNA sequences. Mycol Res\u003cem\u003e\u0026nbsp;\u003c/em\u003e106(9):1043\u0026ndash;1059 https://doi.org/10.1017/S095375620200641X\u003c/li\u003e\n \u003cli\u003eIvaldi P, Maurice JP, Suberbielle N, Domergue P, Normand A-C (2023) \u003cem\u003eClitocella blancii\u003c/em\u003e, un champignon m\u0026eacute;diterran\u0026eacute;en. Bull F\u0026eacute;d Assoc Mycol M\u0026eacute;diterr 64:17\u0026ndash;31 \u0026nbsp;\u003c/li\u003e\n \u003cli\u003eJian S-P, Bau T, Zhu X-T, Deng W-Q, Yang Z-L, Zhao Z-W\u003cem\u003e\u0026nbsp;\u003c/em\u003e(2020a) \u003cem\u003eClitopilus, Clitocella,\u0026nbsp;\u003c/em\u003eand \u003cem\u003eClitopilopsis\u0026nbsp;\u003c/em\u003ein China. Mycologia\u003cem\u003e\u0026nbsp;\u003c/em\u003e112(2):371\u0026ndash;399 https://doi.org/10.1080/00275514.2019.1703089\u003c/li\u003e\n \u003cli\u003eJian S-P, Karadelev M, Wang P-M, Deng W-Q, Yang Z-L (2020b) \u003cem\u003eClitopilus abprunulus\u003c/em\u003e, a new species from North Macedonia with notes on \u003cem\u003eC\u003c/em\u003e. \u003cem\u003eravus\u003c/em\u003e and pleuromutilin producing taxa. Mycol Prog 19:805\u0026ndash;816 https://doi.org/10.1007/s11557-020-01603-6\u003c/li\u003e\n \u003cli\u003eKalichman J, Kirk PM, Matheny PB (2020) A compendium of generic names of agarics and \u003cem\u003eAgaricales\u003c/em\u003e. Taxon\u003cem\u003e\u0026nbsp;\u003c/em\u003e69(3):425\u0026ndash;447 \u0026nbsp;https://doi.org/10.1002/tax.12240\u003c/li\u003e\n \u003cli\u003eKarstedt F, Capelari M, Baroni TJ, Largent DL, Bergemann SE (2019) Phylogenetic and morphological analyses of species of the \u003cem\u003eEntolomataceae\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAgaricales, Basidiomycota\u003c/em\u003e) with cuboid basidiospores. \u003cem\u003ePhytotaxa\u0026nbsp;\u003c/em\u003e391(1):1\u0026ndash;27 \u0026nbsp;https://doi.org/10.11646/phytotaxa.391.1.1\u003c/li\u003e\n \u003cli\u003eKim CS, Jo JW, Kwag YN, Sung GH, Lee SG, Kim SY, Shin CH, Han SK\u003cem\u003e\u0026nbsp;\u003c/em\u003e(2015) Mushroom Flora of Ulleung-gun and a newly recorded \u003cem\u003eBovista\u0026nbsp;\u003c/em\u003especies in the Republic of Korea. 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Nat Ecol Evol\u003cem\u003e\u0026nbsp;\u003c/em\u003e3(4):668\u0026ndash;678 https://doi.org/10.1038/s41559-019-0834-1\u003c/li\u003e\n \u003cli\u003eVilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several \u003cem\u003eCryptococcus\u0026nbsp;\u003c/em\u003especies. J Bacteriol\u003cem\u003e\u0026nbsp;\u003c/em\u003e172(8):4238\u0026ndash;4246 https://doi.org/10.1128/jb.172.8.4238-4246.1990\u003c/li\u003e\n \u003cli\u003eVizzini A, Musumeci E, Ercole E, Contu M (2011) \u003cem\u003eClitopilus chrischonensis\u0026nbsp;\u003c/em\u003esp. nov. (\u003cem\u003eAgaricales, Entolomataceae\u003c/em\u003e), a striking new species from Switzerland. Nova Hedwigia\u003cem\u003e\u0026nbsp;\u003c/em\u003e92(3\u0026ndash;4):425\u0026ndash;434 https://doi.org/10.1127/0029-5035/2011/0092-0425\u003c/li\u003e\n \u003cli\u003eVizzini A, Ferrari RJ, Ercole E, Fellin A (2018) A new species of \u003cem\u003eRhodocybe\u003c/em\u003e sect. \u003cem\u003eRufobrunnea\u003c/em\u003e (\u003cem\u003eEntolomataceae\u003c/em\u003e, \u003cem\u003eAgaricales\u003c/em\u003e) from Italy. MycoKeys 36: 21\u0026ndash;33 https://doi.org/10.3897/mycokeys.36.27094\u003c/li\u003e\n \u003cli\u003eVizzini A, Consiglio G, Marchetti M (2023) Overview of the European species of the genus \u003cem\u003eClitocella\u0026nbsp;\u003c/em\u003e(\u003cem\u003eEntolomataceae, Agaricales\u003c/em\u003e) with notes on extralimital taxa. Persoonia\u003cem\u003e\u0026nbsp;\u003c/em\u003e50:123\u0026ndash;157 \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; https://doi.org/10.3767/persoonia.2023.50.04\u003c/li\u003e\n \u003cli\u003eVizzini A, Alvarado P, Consiglio G, Angelini C, Marchetti M (2024) [2023] \u003cem\u003eLulesia\u003c/em\u003e Singer (1970), an older name for \u003cem\u003eClitocella\u003c/em\u003e Kluting, T.J. Baroni \u0026amp; Bergemann (2014, \u003cem\u003eEntolomataceae\u003c/em\u003e). RMR, Boll Assoc Micol Ecol Romana numero speciale (fuori serie) 2023 (3):3\u0026ndash;23 https://doi.org/10.57624/AMER.2023.18\u003c/li\u003e\n \u003cli\u003evon Bonsdorff T, Kyt\u0026ouml;vuori I, Vauras J, Huhtinen S, Halme P, R\u0026auml;m\u0026auml; T, Kosonen L, Jakobsson S (2014) Sienet ja metsien luontoarvot. Norrlinia 27:1\u0026ndash;272\u0026nbsp;\u003c/li\u003e\n \u003cli\u003eXiao Y-Q, Xu Y-D, Chen Z-H, He Z-M (2024) \u003cem\u003eLulesia umbrinomarginata\u003c/em\u003e (\u003cem\u003eEntolomataceae\u003c/em\u003e, \u003cem\u003eAgaricales\u003c/em\u003e), a newly discovered species from Southern China. Phytotaxa 650(1):060\u0026ndash;072 \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;https://doi.org/10.11646/phytotaxa.650.1.5\u003c/li\u003e\n \u003cli\u003eWhite TJ, Bruns TD, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In Innis MA, Gelfand DH, Sninsky J, White TJ (eds) PCR protocols: a guide to methods and applications. Academic Press, London, pp 315\u0026ndash;322\u003c/li\u003e\n \u003cli\u003eZhao R-L, Li G-J, S\u0026aacute;nchez-Ram\u0026iacute;rez S, Stata M, Yang Z-L, Wu G, Dai Y-C, He S-H, Cui B-K, Zhou J-L, Wu F, He M-Q, Moncalvo J-M, Hyde KD (2017) A six-gene phylogenetic overview of \u003cem\u003eBasidiomycota\u0026nbsp;\u003c/em\u003eand allied phyla with estimated divergence times of higher taxa and a phyloproteomics perspective. Fungal Divers\u003cem\u003e\u0026nbsp;\u003c/em\u003e84(1):43\u0026ndash;74 \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;https://doi.org/10.1007/s13225-017-0381-5\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1 and 2 are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"mycological-progress","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mypr","sideBox":"Learn more about [Mycological Progress](https://www.springer.com/journal/11557)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/mypr/default.aspx","title":"Mycological Progress","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Basidiomycota, Agaricomycetes, Tricholomatineae, Clitocella, Rhodocybe, cryptic species, multigene analysis, taxonomy","lastPublishedDoi":"10.21203/rs.3.rs-7232140/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7232140/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cem\u003eLulesia fallacioides\u003c/em\u003e (from Finland) and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eparvifallax\u003c/em\u003e (from Switzerland) are described as new species, based on both a morphological and molecular approach, in the \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e complex (\u003cem\u003eLulesia\u003c/em\u003e subgen. \u003cem\u003eParaclitopilus\u003c/em\u003e), a morphology-delimited aggregate of species sharing white basidiomes and a bitter context. Compared to \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, the former differs mainly by wider spores and a gelatinized pileipellis and the latter by smaller basidiomes and shorter spores. The recently described \u003cem\u003eClitocella neofallax\u003c/em\u003e from China is here combined in \u003cem\u003eLulesia\u003c/em\u003e. New collections of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003ecolorata\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003esolaris\u003c/em\u003e made in Italy and Finland have allowed us to better understand the distinctive features and/or to extend the distribution area of ​​these recently established species. The presence of \u003cem\u003eL\u003c/em\u003e. \u003cem\u003efallax\u003c/em\u003e, \u003cem\u003eL\u003c/em\u003e. \u003cem\u003emundula\u003c/em\u003e and \u003cem\u003eL\u003c/em\u003e. \u003cem\u003eobscura\u003c/em\u003e in Finland is here molecularly confirmed for the first time. Finally, a full morphological description of \u003cem\u003eL. alachuana\u003c/em\u003e, a North American species so far described from Florida, is provided based on ancient (holotype included) and recent collections.\u003c/p\u003e","manuscriptTitle":"Two new species in the Lulesia fallax complex (Entolomataceae, Agaricales) from Europe (Fennoscandia and Switzerland), Lulesia neofallax comb. nov., new records of recently described species of Lulesia, and notes on Clitocybe alachuana","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-03 16:23:52","doi":"10.21203/rs.3.rs-7232140/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-08-27T04:59:09+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-08-27T02:32:57+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Mycological Progress","date":"2025-08-22T19:03:47+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-08-11T06:06:44+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mycological Progress","date":"2025-08-07T16:17:30+00:00","index":"","fulltext":""},{"type":"decision","content":"Minor Revisions Needed","date":"2025-08-07T07:25:15+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"mycological-progress","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mypr","sideBox":"Learn more about [Mycological Progress](https://www.springer.com/journal/11557)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/mypr/default.aspx","title":"Mycological Progress","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6cc985b3-e8cb-46d1-a67c-59bdd9e41d43","owner":[],"postedDate":"September 3rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-11-24T16:12:02+00:00","versionOfRecord":{"articleIdentity":"rs-7232140","link":"https://doi.org/10.1007/s11557-025-02100-4","journal":{"identity":"mycological-progress","isVorOnly":false,"title":"Mycological Progress"},"publishedOn":"2025-11-21 15:57:07","publishedOnDateReadable":"November 21st, 2025"},"versionCreatedAt":"2025-09-03 16:23:52","video":"","vorDoi":"10.1007/s11557-025-02100-4","vorDoiUrl":"https://doi.org/10.1007/s11557-025-02100-4","workflowStages":[]},"version":"v1","identity":"rs-7232140","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7232140","identity":"rs-7232140","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}