Amanita species within section Caesareae from the highlands of Laos

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Caesareae , collected in Xieng Khouang and Vientiane provinces in Laos from highland forests dominated by members of Fagaceae and Pinaceae , are studied based on morphological characters and analyses of ITS sequences. Two species, namely A. fulvocaesarea and A. chepangianoides are reported as new, both found in association with Fagaceae , whereas A. pseudoprinceps and A. subhemibapha , recently described from China and A. rubromarginata from Japan, are confirmed from Laos and found in forests with Fagaceae and Pinaceae . The phylogenetic analyses to recognize the Amanita species showed that the golden-brown to greyish orange A. fulvocaesarea and the dominantly white A. chepangianoides cluster with A. aporema, A. chepangiana, A. egregia , A. princeps and A. pseudoprinceps. A dichotomous key to identified and likely occurring Amanita species in Laos within sect. Caesareae is also provided. Amanitaceae taxonomy South–East Asia Fagaceae Pinaceae 2 new taxa Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction The genus Amanita Pers. (1797: 65) ( Amanitaceae, Agaricales) is widely occurring in temperate, subtropical, and tropical regions and contains approximately 1,200 binomials of which some 650 are currently accepted species (Yang et al. 2018 ; Tulloss and Yang 2024 ). Mycorrhizal tree partners are foremost with Fagaceae , Pinaceae , Dipterocarpaceae and Betulaceae (Tulloss and Yang 2024 ), all with members in Laos (Newman et al. 2007 ). Currently, three subgenera are accepted within Amanita - Lepidella, Amanitina and Amanita - with subgenus Amanita having four sections: Amanita, Amarrendiae, Caesareae , and Vaginatae (Cui et al. 2018 ). This study is limited to identified Lao specimens within sect. Caesareae . This section, when agaricoid, is characterized by fruitbodies with a striate-sulcate pileus margin, truncate lamellulae, annulate stipes, a non-bulbous stipe base with a saccate volva, inamyloid basidiospores and presence of clamps (Cui et al. 2018 ). The Amanitaceae.org website lists 101 binomials within sect. Caesareae of which 71 have a detailed description (Tulloss and Yang 2024 ). Sect. Caesareae has an almost cosmopolitan distribution with many species found in South and East Asia and in recent years several new species have been published from this region foremost from China, where for example six new species were added by Cui et al. ( 2018 ). Laos lies within the Indo-Burma Biodiversity Hotspot (Myers 1988 ) and boasts a relative high forest coverage of 56%, spanning from dipterocarp dominated forests at lower altitudes to broadleaved-coniferous and evergreen forests at higher altitudes (MAF 2021). These forest types accommodate numerous ectomycorrhizal fungi, including many Amanita species (Læssøe et al. 2018 ). Patouillard ( 1928 ) was probably the first to publish on Laotian macrofungi among mainly aphyllophoroid species from Vietnam. Another early publication mentioning macrofungi in Laos was the Flora of Laos by Vidal ( 1958 ), a French botanist, who included 33 macrofungal species. Much later, this was followed-up by a non-timber forest products handbook (NAFRI 2007), which included 29 common, edible and medicinal fungal species. Recent surveys by mycologists have been carried out at biodiversity conservation areas (Sysouphanthong et al. 2017 ; Lee et al. 2018 ; Hong et al. 2019 ; Lee et al. 2021 ), at local community forests (Pedersen and Thammavong 2014 ) and at local markets (Łuczaj et al. 2021 ). Macrofungal species described based on material from Laos are rather few, but increased in recent years and includes Phellinus stratosus Pat. (Patouillard 1928 ), Cyptotrama angustispora Zhu L. Yang & J. Qin and C. glabra Zhu L. Yang & J. Qin (Qin and Yang 2016 ), Leucoagaricus houaynhangensis Sysouphanthong (Sysouphanthong et al. 2018 ), Ganoderma nasalaense Hapuar., Pheng. et K.D. Hyde (Hapuarachchi et al. 2019 ), Hebeloma parvisporum O. S. Pedersen, Læssøe, Beker & Eberh. (Eberhardt et al. 2020 ), Cystolepiota pyramidalis Sysouph. & Thongkl. and C. rhodella Sysouph. & Thongkl. (Sysouphanthong et al. 2022 ), Nigrocarnea radicata O. S. Pedersen & Læssøe (van de Peppel et al. 2022 ), Leucoagaricus flavus Sysouph. & Thongkl. and L. griseosquamosus Sysouph. & Thongkl. (Sysouphanthong and Thongklang 2022 ), and L. laosensis Sysouph. (Manawasinghe et al. 2022 ). Læssøe et al. ( 2018 ) published the first comprehensive introductory field guide on edible, poisonous and medicinal mushrooms of Northern Laos including 24 Amanita species of which three were from sect. Caesareae . The authors noted that many specimens need supporting molecular analyses to confirm their identity or make the description of new species possible. This study is the first of its kind from Laos, with the aim to document Lao species of genus Amanita sect. Caesareae , using morphological characters, molecular phylogenetic evidence and ecological data. Materials and methods Basidiomata were collected during the monsoon season from May to September from rich mixed forests, mainly dominated by Fagaceae and Pinaceae in the highland areas of Xieng Khouang and Vientiane provinces, Laos (Lao People's Democratic Republic). Specimens were deposited in the National Herbarium of Laos (HNL) with duplicates of some collections in C. Index Fungorum ( http://www.IndexFungorum.org ), the International Index of Plant Names ( https://www.ipni.org ) and Amanitaceae.org ( http://www.Amanitaceae.org ) were used as sources for taxonomic names and nomenclature. Macroscopic characters were described from fresh material with field notes and photographs. Pileal sulcation is annotated as the ratio (R) of the length of the striations from the pileal margin to the centre of the pileus. Stipe length is from the volval base. Colour codes follow Kornerup and Wanscher ( 1981 ). Microscopic structures were observed with light microscopy (Brunel SP500 with OMAX 14MP mount microscope camera) from the dried specimens after sectioning and mounting in H 2 O and 3% KOH solution and where necessary stained with Congo Red. Basidiospores were tested for amyloidity with Melzer’s reagent and were measured in 3% KOH. The abbreviation (n/m/p) means n basidiospores measured from m basidiomata from p collections, using a range notation of the form (a–) b–c (–d), the range b–c contains a minimum of 90% of the measured values, extreme values (a, d) are given in parentheses, m and av. indicate average values, ± standard deviation; Q presents the length/width ratio of spores in side view. Description terminology follows Bas ( 1969 ), Tulloss and Rodríguez-Caycedo ( 2011 ) and Tulloss and Yang ( 2024 ). Sizes of basidiomata are described as small, medium or large with pileus diameter of 30–50 mm, 50–90 mm, and 90–150 mm, respectively (Bas 1969 ). DNA extraction and sequencing of the ITS1, 5.8S, ITS2 regions were carried out at Tartu University, Estonia and uploaded into PlutoF cloud database (Abarenkov et al. 2010 ), following the procedures referred to and described in Voitk et al. ( 2020 ). Lao sequences were blasted in Unite website, https://unite.ut.ee/analysis.php (Kõljalg et al. 2020 and Abarenkov et al. 2023 ) and at NCBI website, http://www.ncbi.nlm.nih.gov/blast and related sequences (supplementary table 1) were retrieved for phylogenetic analyses. Priority was given to similar species, as well as to species representing neighbouring clades within sect. Caesareae described in recent publications (Tang et al. 2017 ; Cui et al. 2018 ; Kumla et al. 2023 ). The final ingroup dataset consisted of 38 sequences of which 12 new ITS sequences were from Laos, and A. pseudosychnopyramis from sect. Amanita was selected as outgroup. Supplied by the NGPhylogeny website http://www.phylogeny.fr/index.cgi (Lemoine et al. 2019 ) sequences was aligned with MUSCLE (Edgar 2004 ) and MAFFT v.7.4 (Katoh and Standley 2013 ) and inspected for reverse sequences in BioEdit (Hall 1999 ); poorly aligned positions were excluded applying the BMGE (Block Mapping and Gathering with Entropy) software (Criscuolo and Gribaldo, 2010 ), Maximum Likelihood (ML) phylogenetic analysis were performed by PhyML v.3.3 (Guindon et al. 2010 ) + Smart Model Selection (SMS) (Lefort et al. 2017 ) and by Bayesian Interference (BI) analysis by MrBayes v. 3.2.7 (Ronquist et al. 2012 ). The final trees were edited and visualized using Interactive Tree Of Life (iTOL) v.6 (Letunic and Bork 2021 ) and Microsoft Office. The BMGE selected 532 and removed 998 characters. MrBayes for BI was performed with 1.000.000 generations and otherwise default settings. The ML tree was examined by HKY85 substitution model with 1,000 replicates, otherwise default settings resulting in the log-likelihood tree value of -3698.83699 for the best scoring tree. The phylogram topology derived from BI and PhyML were similar and only the ML tree is shown in Fig. 1 . In comparison, while running ML in MEGA11 (Tamura et al. 2021 ) a similar topology and bootstrap values were observed. A branch was considered strongly supported if it had ML bootstrap support ≥ 85% and posterior probability (PP) ≥ 0.95. Dataset and final alignments for A. fulvocaesarea and A. chepangianoides have been deposited in TreeBase (No. xxxxxxx/xxxxxxx and xxxxxxxxxxxx/xxxxxxx, respectively. Results Taxonomy Etymology Epithet refers to the similarity to Amanita chepangiana Tulloss & Bhandary. Holotype : Laos. Xieng Khouang province, Phoukhout district, Ngodphae village, Na Phouak hamlet, 19°45'43.8"N, 103°15'22.5"E, 1385 m asl., in broadleaved forest with Quercus spp. Castanea spp. and Lithocarpus spp., 23 May 2016, O. Sparre Pedersen and T. Læssøe, OSP20160523–013, HNL501136, as A. ? chepangiana in Læssøe et al. ( 2018 ), isotype C-F-169624 (C), GenBank: ITS = xxxxxxxxx Diagnosis Amanita chepangianoides is characterized by white basidiomata except the alutaceous-cream centre of the pileus, a relative short pileal margin striation (0.25 R), a snake-skin pattern on the stipe and globose to subglobose spores (8.5–10.5 × 8.5–9.5 µm) and presence of clamps. Description Basidiomata (Fig. 2 a, f) medium to large. Pileus 90–130 mm in diam, initially almost spheric then convex and centrally depressed, slightly viscid, cream (4A2–3) to alutaceous cream (5B4–7B4) at centre and whitish (5A1–2) towards margin; velar remnants absent; margin striate (0.25 R), non-appendiculate; trama white (1A1), unchanging. Lamellae free or almost so, plentiful (approximately 100 reaching stripe), white (1A1), edge fimbriate, concolorous when fresh and when dried; lamellulae truncate, few of 2–3 lengths. Stipe 160–200 × 10–18 mm, white (1A1), cylindrical, slightly tapering upwards, basal bulb absent, covered by white (1A1) squamules, mostly in a snake-skin pattern; context white (1A1), unchanging; volva large, ca. 45 mm high and 35 mm wide, flaring, saccate, white (1A1). Annulus apical, membranous, white (1A1), upper surface sometimes weakly striate, hanging, fugacious. Taste not recorded. Smell pleasant, not prominent. Spore deposit not observed. Lamellar trama (Fig. 2 b). Mediostratum 20–40 µm wide, composed of fairly abundant ellipsoid to fusiform to cylindrical, thin-walled inflated cells, 25–72 × 17–32 µm; filamentous hyphae abundant, 1–2 µm wide; lateral stratum 37–55 µm wide, composed of ellipsoid to clavate inflated cells (15–40 × 10–15 µm), filamentous hyphae scarce, 1–2 my wide. Subhymenium (Fig. 2 d) 25–42 µm thick composed of 2–3 layers of subglobose, ovoid, ellipsoid to clavate cells, 12–20 × 9–14 µm. Lamellar edge sterile, composed of irregular inflated cells, subglobose to irregular, hyaline, thin-walled, 16–29 × 15–25 µm; filamentous hyphae scattered, 0.5–2 µm wide. Basidia (Fig. 2 d) 22–35 × 8–14 µm, clavate, mainly 2-spored, few 4-spored; sterigmata 3–6 µm long. Basidiospores (Fig. 2 e) [40/1/1] (8–) 8.5–10.5 (–11) × (7.5–) 8.5–9.5 (–10.5) µm (av. 9.6 × 8.8 µm), Q = 1.04–1.14 (Qm = 1.07 ± 0.06), globose to subglobose (few broadly ellipsoid), inamyloid, hyaline, thin-walled, smooth; apiculus 1–2 µm my high and 1–1.5 µm wide. Pileipellis 80–120 µm thick; upper layer 25–70 µm, gelatinized, composed of radically arranged, filamentous hyphae 2–10 µm wide; lower layer 25–50 µm composed of radically arranged filamentous hyphae, 4–8 µm wide, hyaline to yellowish. Interior of volva (Fig. 2 c) composed of longitudinally arranged elements; filamentous hyphae very abundant 1–7 µm wide, few branching; inflated cells scattered to locally abundant, subglobose, broadly ellipsoid, ellipsoid to cylindric 25–62 × 15–45 µm, hyaline to yellowish, thin-walled; gelatinized hyphae locally abundant. Stipe trama composed of longitudinally arranged, mainly cylindrical terminal cells, thin-walled, 30–240 × 14–65 µm; filamentous hyphae scattered, 3–10 µm wide. Annulus composed of interwoven filamentous hyphae and inflated cells; inflated cells rare, subglobose to ellipsoid, 25–55 × 24–35 µm; filamentous hyphae abundant, 2–8 µm wide. Clamps presumably present in all parts, but difficult to observe, seen with certainty on stem hyphae. Habitat Scattered on rich soil in broadleaved highland forests under Fagaceae . Additional specimen examined Laos. Xiang Khouang province, Phoukhout district, Ngodphae village, 19°45'43.3"N, 103°15'23.0"E, 1380 m asl., in broadleaved forest with Quercus spp., Castanea spp. and Lithocarpus spp., 23 May 2016, O. Sparre Pedersen and T. Læssøe, OSP20160523-012 (HNL501150). Discussion Our ITS phylogenetic analysis (Fig. 1 ) places A. chepangianoides in a larger clade containing A. chepangiana Tulloss & Bhandary, A. fulvocaesarea O.S. Pedersen & T. Læssøe (described as new in this study), A. princeps Corner and Bas, A. aporema Boedijn, A. egregia D.A. Reid and A. pseudoprinceps Y.-Y. Cui, Q. Cai & Zhu L. Yang. Morphologically, the similar A. chepangiana differs by sometimes having yellowish to greyish tinges at centre of the pileus and by subglobose to ellipsoid basidiospores (9.5–12.5 × 8.5–11.5 µm, Q = 1.07–1.28, Qm = 1.17 ± 0.06) (Tulloss and Bhandary 1992 ; Cui et al. 2018 ). Amanita alboumbelliformis Y.-Y. Cui, Q. Cai & Zhu L. Yang is also similar by sometimes having a cream pileus centre, but it differs by having a longer pilea striation (0.3–0.4 R), larger spores (9–11 × 9–11 µm, Q = 1.0–1.13, Qm = 1.06 ± 0.03), yellowish tinges on the aging annulus and a greyish to brownish volva (Cui et al. 2018 ). Amanita egregia has white basidiomata, but differs by having larger pilei and stipes (up to 200 mm), broadly ellipsoid spores and association with Eucalyptus in Australia (Reid 1978 ). Amanita fulvocaesarea O. S. Pedersen & T. Læssøe, sp. nov. (Fig. 3 ) MycoBank MB xxxxx. Holotype : Laos. Xiang Khouang province, Phoukhout district, Ngodphae village, Na Phouang hamlet 19°45'11.9"N, 103°15'29.1"E, 1135 m asl., on soil with Quercus spp., Castanopsis spp. and Lithocarpus spp., 11 May 2015, O. Sparre Pedersen and T. Læssøe, OSP20150511–007, HNL500511, as A. princeps in Læssøe et al. ( 2018 ), isotype C-F-169623 (C), GenBank Acc. No.: ITS xxxxxxxxx. Diagnosis : Amanita fulvocaesarea is characterized by the initial greyish-orange colours followed by golden brown hues, an umbonate pileus, distinct sulcation (0.3 – 0.4R), an apical annulus, white non-cracking volva, globose to subglobose basidiospores (9–12 x 9–11.5 µm) and presence of clamps. Description Basidiomata medium to large (Fig. 3 a, f). Pileus up to 140 mm in diameter, first ovoid then convex to applanate with small broad umbo, dry to slightly viscid, centre first greyish-orange (5B4), with age golden brown (5D6) with tinges of greyish-ochraceous (5–8B2) towards margin; veil remnants absent; margin sulcate (0.3–0.4 R), non-appendiculate; trama white (1A1), unchanging. Lamellae free or almost so, whitish (1A1), when dry brownish-orange (6C5), appr. 100 reaching the stipe; lamellulae few of 2–3 lengths, truncate; edge fimbriate, sterile, concolorous, when dried whitish-cream (4A1–2). Stipe 150–200 ×12–20 mm, white (1A1) smooth to slightly fibrous, tapering towards apex, sometimes almost cylindrical; context white (1A1), hollow; basal bulb absent; volva 30–60 mm high and 20–40 mm wide, saccate, thick, white (1A1), not cracking, flaring. Annulus white (1A1) on both sides, apical, big, pendent, but rather fugacious; upper surface non-striate. Taste not recorded. Odour indistinctive. Spore deposit not observed. Lamellar trama (Fig. 3 b) bilateral. Mediostratum 42–63 µm wide, interwoven, inflated cells thin-walled, hyaline, scattered, fusiform, ellipsoid to clavate 18–180 × 7–27 µm; filamentous hyphae abundant, 1–5 µm wide; lateral stratum 36–64 µm wide with abundant clavate to cylindrical, inflated cells 23–38 × 5–11 µm; filamentous hyphae scarce, 1–2 my wide. S ubhymenium (Fig. 3 d) 14–34 µm thick, composed of 2–3 layers of subglobose, ellipsoid to irregular cells, 8–21 × 6–15 µm. Lamellar edge sterile composed of scattered subglobose, ellipsoid to cylindric, inflated, hyaline, thin-walled cells, 16–36 × 11–18 µm and scattered filamentous, 1–2.5 µm wide. Basidia (Fig. 3 d) 42–50 × 15–22 µm, clavate, 2- (few) and 4-spored, sterigmata 3–6 µm long. Basidiospores (Fig. 3 e) [30/1/1] 9–12 (–12.5) × 9–11.5 (–12) µm (av. 11.5 × 11 µm), Q = 1.01–1.08, (Qm 1.04 ± 0.03); globose to subglobose, inamyloid, hyaline, thin-walled, smooth; apiculus 1.5–2 µm high and 1–2 µm wide. Pileipellis 135–200 µm thick; upper layer 40–140 µm, gelatinized, composed of radially arranged to interwoven hyphae, 4–11 µm wide, hyaline to yellowish-brown; lower layer 30–75 µm thick, composed of interwoven filamentous hyphae 1.5–3.5 µm wide, hyaline to yellowish. Volva dominantly composed of filamentous hyphae; interior surface (Fig. 3 c) composed of longitudinally orientated filamentous hyphae, 2–4 µm wide, hyaline, thin-walled; outer surface of similar structure, but with gelatinized filamentous hyphae 2–6 µm wide; inflated cells scattered, subglobose, fusiform to ellipsoid, 15– 90×10–50 µm, hyaline, thin-walled. Stipe trama composed of longitudinally arranged, inflated, cylindrical to slightly clavate, hyaline, thin-walled terminal cells, 67–215 × 31–45 µm; filamentous hyphae abundant, 3–5 µm wide. Annulus composed of interwoven elements: outer part with inflated cells rare to scattered, subglobose to ellipsoid, 15–34 × 13–23 µm, hyaline, thin-walled; filamentous hyphae very abundant 1–5 µm wide, hyaline, thin-walled; inner part with radially arranged inflated cells, scattered to fairly abundant, hyaline, subglobose to elongate, 20–44 × 8–24 µm; filamentous hyphae abundant, 1–6 µm wide, hyaline, yellow to yellowish-brown, thin-walled. Clamps presumably present in all parts, but difficult to observe, seen with certainty on pileipellis hyphae and on stem hyphae. Habitat and distribution On rich soil in broadleaved forests with members of Fagaceae (Laos) Fagaceae and Pinaceae (China). Known from Laos (this study) and China (non-described). Additional specimen examined Laos. Xiang Khouang province, Phoukhout district, Ngodphae village, 19°45'43.3"N, 103°15'23.0"E, 1381 m asl., in broadleaved forest with Quercus spp., Castanopsis spp. and Lithocarpus spp., 23 May 2016, O. Sparre Pedersen and T. Læssøe, OSP20160523–009 (HNL501130). China. Yunan province, Puer, ca. 1400 m asl., in a forest with Lithocarpus and Pinus , 6 July 2015, Qing Cai 1349 (KUN-HKAS91972). The analyses of the Chinese taxa are based on Genbank ITS sequences, only. Discussion Our phylogenetic analysis (Fig. 1 ) showed that three Amanita specimens: HKAS91972 (ITS = MH508608), HKAS101390 (ITS = MH508603) and HKAS91096 (ITS = MH508607) from China, clustered with A. fulvocaesarea with strong ML/PP support. These specimens were briefly mentioned by Cui et al. ( 2018 ), but not assigned to any described taxon, due to insufficient morphological observations. For related species in the clade please see the commentary section of A. chepangianoides . Morphologically A. fulvocaesarea is similar to A. pseudoprinceps and A. princeps . The former differs by a shorter pileal striation (0.2–0.3 R) and subglobose to broadly ellipsoid spores (Q = 1.02–1.23, Qm = 1.12 ± 0.04) (Cui et al. 2018 ) and in this study (Q = 1.02–1.25, Qm = 1.12 ± 0.08). The latter species differs by a larger (up to 200 mm) brownish to yellow pileus, shorter pileal striation (0.2–0.25 R), a volval outer surface that often cracks and peels into thin brownish patches (Corner and Bas 1962 ; Yang 2015 ; Tang et al. 2017 ) and the association with dipterocarps (Malaysia) (Tang et al. 2017 ). Amanita aporema has some similarities, but it differs by relatively smaller basidiomata and darker coloured pilei (brown to reddish-brown, ca. 60–100 mm in diam.) and longer striations, ca. 0.5 R (Tang et al. 2017 ; Cui et al. 2018 ). Description Basidiomata (Fig. 4 a, d-f) medium sized to large. Pileus 60–140 mm in diam, initially hemispheric, then convex to applanate, slightly viscid, olive grey (1E2) to yellowish-brown (5D–E8), darker towards centre and white to yellowish-cream (3–4A3) to white (1A1) at margin; velar remnants absent; margin striate (0.2–0.25 R), non-appendiculate; trama white (1A1), unchanging. Lamellae free or almost so, white (1A1) to whitish-cream (4A1–2), plentiful (> 100 reaching stipe), up to 20 mm wide; lamellulae truncate, few of 5–6 lengths. Stipe 70–150 × 10–20 mm, cylindrical, basal bulb absent, white (1A1) with slightly darker fibrils; context white (1A1), unchanging, hollow; volva large, saccate, 30–60 mm high and 20–40 mm wide, white (1A1), sometimes brownish (2B2-4) patches. Annulus (Fig. 4 f) apical, skirt-like, membranous, 10–20 mm high, white (1A1), partly striate on upper surface. Taste not recorded. Smell insignificant. S pore deposit not observed. Basidia (Fig. 4 b) clavate, 28–54 × 10–17 µm, 4-spored, sterigmata 1.5–7 µm long. Basidiospores (Fig. 4 c) [120/3/3] (8.5–) 9.5–14.5 (–18.5) × (7.5–) 8–12.5 (–13.5) µm (av. 12 × 10.5 µm), Q = 1.02–1.25 (Qm = 1.12 ± 0.08), subglobose to broadly ellipsoid, very few globose, inamyloid, hyaline, thin-walled, smooth; apiculus 1.5–2 µm high and 1–1.5 µm wide. Pileipellis 95–150 my thick; upper layer composed of radically arranged gelatinized filamentous hyphae, 1–2 µm wide; lower layer composed of radically arranged, hyaline, filamentous hyphae, 4–8 my wide. Interior of volva composed of filamentous hyphae 1.5–8 µm wide; inflated cells scattered, ellipsoid, hyaline, hyaline, thin-walled, scattered to locally abundant, 22–55 × 13–25 µm. Stipe trama composed of longitudinally arranged, ellipsoid, clavate to cylindric, thin-walled terminal cells, 85–170 × 15–60 µm; filamentous hyphae scattered to abundant, 2–10 µm wide. Clamps not observed. Habitat and distribution Scattered on soil in mixed forests with members of Fagaceae , Dipterocarpaceae and Pinaceae or without the dipterocarp element, at altitudes 800–1125 m, May to August. Known from southwestern China, Nepal, Laos and Thailand. Common at local markets in Laos. Specimens examined Laos. Vientiane province, Thoulakhom district, Vangheua village, 18°20'31.5"N, 102°47'46.6"E 790 m asl., in mixed forest with Pinus kesiya , Dipterocarpus spp. and Fagaceae , 19 May 2012, O. Sparre Pedersen, OSP20120519-001 (HNL500333). Ibid., 18°20'39.7"N, 102°47'36.9"E. 803 m asl., 6 July 2013, O. Sparre Pedersen, OSP20130706-004 (HNL500044). Xieng Khouang province, Pek district, Dong Village, 19°29'39.1"N, 103°16'07.7"E 1123 m asl., in mixed forest including Pinus kesiya , Keteleeria everlyniana and Quercus spp. 22 June 2014, O. Sparre Pedersen, OSP20140622–007 (HNL500154). Discussion Amanita pseudoprinceps was recently described from Yunnan, China, from a Fagaceae dominated mixed forest at 2300 m altitude (Cui et al. 2018 ). The Laotian specimens (HNL500044, HNL500155 and HNL500333) match the type in terms of ecology, pileus and stipe colours, striation on pileus, apical placement of annulus, spore size as well as in our phylogenetic analysis (Fig. 1 ). A minor difference is that Laotian specimens have a hollow stipe in full length, compared to the holotype, which is only hollow in the centre. As discussed above, A. pseudoprinceps can be easily confused with A. princeps , which differs by a broader pileus (100–200 mm), with light brown to yellowish-brown colours and with brownish cracking patches on the outer surface of the volva and association with dipterocarps (Tang et al. 2017 ). It also resembles A. fulvocaesarea (described as new above), which is smaller, has a longer pileal striation (0.3–0.4 R) and dominantly globose spores. The two other species shown in the clade (Fig. 1 ), A. aporema and A. egregia differ from A. pseudoprinceps by a brown to reddish-brown pileus and a white basidioma, respectively (Tang et al. 2017 ; Reid 1978 ). Recent work of Łuczaj et al. ( 2021 ) surveying markets in Luang Prabang, Laos, as well as an ethnobotanical survey in Yunnan, China by Wang et al. ( 2022 ) identified two taxa as A. princeps (WA0000072256: ITS = MT 252585 and HKAS122502: ITS = ON394332, respectively). With strong bootstrap support in our study (Fig. 1 ) these taxa should be referred to as A. pseudoprinceps . See Cui et al. ( 2018 ) for further misidentified specimens. Amanita rubromarginata Har. Takah., Mycoscience 45(6): 372 (2004) (Fig. 5 ) Description Basidiomata (Fig. 5 a, b, f) medium sized. Pileus 45–90 mm in diam, first almost cylindrical, then convex to applanate, centrally depressed, occasionally with broad umbo, dry or almost so, orange red (8A8) to brownish-red (9C–D8) at centre and orange (5A6–7), yellowish-orange (4AB7) to deep yellow (4A8) at margin; velar remnants absent; margin striate (0.4–0.5 R), non-appendiculate, trama yellowish (4A4–5). Lamellae free or almost so, plentiful (ca. 110 reaching stem), yellowish (2A7); edge fimbriate, light orange (5A4) to reddish-orange (7A7); lamellulae truncate, few of 3–4 lengths. Stipe 70–120 × 7–9 mm, cylindrical to slightly tapering upwards, basal bulb absent, hollow, yellow (2A7–3A7) to light yellow (4A4–5), partially covered by orange (5A6) to orange-red (8A8) squamules in a snake-skin pattern; context light yellow (4A4–5); volva thick, large, saccate, 35–40 mm high and 25–30 mm wide, whitish (2A2). Annulus subapical, membranous, hanging, dark orange (5A8) to reddish-orange (7A7), upper surface non-striate, edge dentate. Taste not recorded. Smell indistinct. Spore deposit not observed. Basidia (Fig. 5 c) 9–12 x 26–37 µm clavate, 4-spored; sterigmata 2.5–4.5 µm long; basal septa clamped. Basidiospores (Fig. 5 d) [40/2/2] 6.5–9 (–9.5) × 5.5–7 (–7.5) µm (av. 7.5 × 6.2 µm), Q: 1.13–1.37 (Qm = 1.22 ± 0.7), broadly ellipsoid, few subglobose and few ellipsoid, inamyloid, hyaline, thin-walled, smooth; apiculus small, 0.5 µm high and 1–1.5 µm wide. Pileipellis 80–130 µm thick, 2-layered with radically arranged hyphae; upper layer 30–35 µm wide, filamentous hyphae gelatinized, few branching, thin-walled, hyaline, 1–6 µm wide; lower layer 50–100 µm thick, gelatinized; filamentous hyphae 3–8 µm wide, hyaline to yellowish. Interior of volva composed of longitudinally arranged elements: filamentous hyphae 2–7,5 µm wide, hyaline to yellowish; inflated cells scattered to abundant ellipsoid to elongate 37–54 × 23–40 µm, hyaline to yellowish, thin-walled. Stipe trama composed of longitudinally arranged, mainly cylindrical, terminal cells, 100–150 × 15–30 µm; filamentous hyphae scattered, thin-walled, 3–7 µm wide. Clamps not observed with certainty. Habitat and distribution Scattered, on soil in mixed highland forests with species of Pinaceae and Fagaceae . Known from China, Loas (this study) Japan, Malaysia and Thailand. Specimens examined Laos. Xieng Khouang province, Pek district, Dong village, 19°29'37.8"N, 103°16'11.2"E1121 m asl., in mixed forest with Quercus spp. , Pinus kesiya , and Keteleeria everlyniana , 14 August 2015, O. Sparre Pedersen and T. Læssøe, OSP20150814-016 (HNL500859); ibid., 19°29'39.8"N, 103°16'05.8"E, 1129 m asl., 16 August 2022, O. Sparre Pedersen and W. Sibounheuang, OSP20220816-005 (HNL502458). Discussion Amanita rubromarginata was described from broad-leaved forests in southwestern islands of Japan (Takahashi 2004 ). It is distinctive by the long striation, i.e. 0.5 R in Cui et al. ( 2018 ) (in this study 0.4–0.5 R), the reddish-orange lamellar edge, dark orange to reddish-orange membranous annulus, thick white volva as well as the association with Fagaceae . Notably, we have not seen a greyish-yellow discoloration of aged specimens as shown in Terashima et al. ( 2016 ). Similarly, this feature was similarly not recorded in the description of this species from China (Cui et al. 2018 ) and from Thailand (Thongbai et al. 2016 ; Kumla et al. 2023 ). Amanita rubromarginata somewhat resembles A. malayensis L.P. Tang, Zhu L. Yang & S.S. Lee and A. javanica (Corner & Bas) T. Oda, C. Tanaka & Tsuda. The former differs by a yellowish to yellowish-orange pileus, concolorous lamellae edge, and the latter differs by less reddish to orange tinges on pileus and annulus. Both of these species are recorded in association with members of Dipterocarpaceae (Tang et al. 2017 ; Cui et al. 2018 , Tulloss and Yang 2024 ). Amanita subhemibapha Zhu L. Yang, Y.-Y. Cui & Q. Cai (see below) differs by lacking the dark lamellae edge and yellowish zone in lower pileipellis. In our phylogenetic tree (Fig. 1 ) the three lao specimens clustered with strong support the A. rubromarginata specimens from China and Thailand. Description Basidiomata (Fig. 6a, b, e) medium to large sized. Pileus up to 100 mm in diam, initially almost cylindrical, then applanate to slightly depressed, not umbonate, dry or almost so, centre brownish-orange (7C6–7) when young then reddish-orange (7A8), orange (5A6, 6A8), at margin orange-yellow (4A8); veil remnants absent; margin striate, 0.2–0.4 R; non-appendiculate; trama white (1A1), unchanging; lower pileipellis with a pale to light yellow (3A3-5) zone, 0.7–0.9 mm wide. Lamellae free or almost so, yellowish-white (3A2) to light yellow (4A5), 9–11 mm wide, plentiful (nearly 100 reaching stipe); edge light to pale yellow (3A3–4); lamellulae truncate, l–2 shorter of various lengths between full-length lamellae. Stipe 80–110 × 8–10 mm, cylindrical, hollow, yellowish-white (3A2), partly to fully covered with reddish-orange (7A8) to dark orange (5A8) fibrils; volva saccate, thin, 35–60 mm high and 30–40 mm wide, white (1A1). Annulus subapical, membranous, hanging, yellowish-white (3A2) to dark orange (5A8) upper/lower concolorous; upper surface non-striate. Taste not recorded. Smell indistinct. Spore deposit white (1A1). Basidia (Fig. 6c) 27–55 × 10–16 µm, clavate, 4-spored; sterigmata 2–5 µm long. Basidiospores (Fig. 6d) [80/2/2] 7.5–10 (–10.5) × (6–) 6.5–8 (–8.5) µm (av. 8.5 × 7 µm), Q: 1.05–1.39 (Qm 1.23 ± 0.08) broadly ellipsoid to ellipsoid, few subglobose, inamyloid, hyaline, thin-walled, smooth; apiculus 0.5–1 µm high and 1–1.5 µm wide. Pileipellis 120–175 µm thick; upper layer 50–100 µm thick, gelatinized, composed of radially to interwoven, thin-walled, hyaline, filamentous hyphae, 2–3 µm wide; lower layer 90–120 µm thick, composed of radially to interwoven filamentous hyphae, 4–15 µm wide, hyaline. Interior of volval remnants composed of longitudinally to interwoven arranged elements; filamentous hyphae abundant, 1.5–10 µm wide, hyaline, thin-walled; inflated cells scattered to abundant, broadly ellipsoid, elongate to narrowly cylindrical, hyaline, thin-walled, 38–125 × 30–60 µm. Stipe trama composed of longitudinally arranged, mainly cylindrical, terminal cells, thin-walled, hyaline, 105–200 × 24–46 µm; filamentous hyphae scattered, 1.5–6 µm wide. Habitat and distribution Scattered on soil in mixed highland forests at altitudes of ca 1150 m, dominated by members of Pinaceae and Fagaceae . Know from Southwestern China, Laos (this study) and northern Thailand. Specimens examined Laos. Xieng Khouang province, Pek district, Boa village, 19°29'47.3"N, 103°17'39.1"E, 1137 m asl., in mixed forest with Castanopsis spp., Quercus spp. , few Pinus kesiya and few Keteleeria everlyniana , 6 August 2017, O. Sparre Pedersen and W. Sibounheuang, OSP20170806-005 (HNL501890); ibid., Dong village, 19°29'32.7"N, 103°16'06.2"E, 1141 m asl., in mixed forest with Quercus spp. , few Pinus kesiya and few Keteleeria everlyniana , 31 July 2018, O. Sparre Pedersen, OSP20180731–003 (HNL502059), as A . cf. hemibapha in Læssøe et al. ( 2018 ). Discussion Amanita subhemibapha was recently described from Yunnan, southwestern China, in association with Fagaceae and Pinaceae and recognized by the combination of a non-umbonate orange centre of the pileus, striation 0.25–0.35 R, white to cream lamellae, yellowish lamellae edges and broadly ellipsoid to ellipsoid spores (Cui et al. 2018 ). The Laotian specimens are slightly darker in the yellow and orange colours; the veil remnants on the stipe are more distinct, and the yellow zone in lower pileipellis is also more pronounced. Amanita subhemibapha somewhat resembles A. hemibapha (Berk. & Broome) Sacc., A. malayensis, A. kitamagotake N. Endo & A. Yamada, A. rubromarginata and A. javanica. Amanita hemibapha differs by having a darker red pileus centre, an orange margin and narrower spores (5.5–6.5 µm) (Vrinda et al 2005 ; Cui et al. 2018 ) to more yellowish pileus margin and spore width of 5.5–7 µm (Kumla et al. 2023 ). Amanita rubromarginata and A. javanica differ by having longer pileal striation (0.4–0.5 R) and reddish lamellae edges; A. kitamagotake differs by having uniformly yellow colours on pileus and stipe surface (Yang 2015 ; Endo et al. 2017 ; Cui et al. 2018 )d malayensis differs by having a whitish-cream stipe and an association with dipterocarps (Tang et al. 2017 ). Our phylogenetic tree (Fig. 1 ) indicates that A. subhemibapha is closely related to A. malayensis A. similis Boedijn and A. fuscoflava , Zhu L. Yang, Yang-Yang Cui & Qing Cai. The former differences to A. subhemibapha are discussed above and under the discussion heading of A. rubromarginata; A. similis differs by a longer pileal striation (0.33–0.6 R) and a greyish brown to olivaceous brown pileal center (Boedijn ( 1951 ; Corner and Bas 1962 ); and A. fuscoflava also differs by a longer pilar striation (0.5–0.6 R) and a dark brown pilar center (Cui et al. 2018 ). Key to Amanita sect. Caesareae known from and likely to occur in Laos Stipe white to cream with fibrils of same colours 2 Stipe yellowish-white to yellowish, typically with darker fibrils 8 2. Basidiomata entirely white or cream to alutaceous-cream at centre of pileus 3 Basidiomata with yellow-greyish, cream, brown to brownish centre of pileus 5 3. Striation on pileus 0.3–0.4 R; annulus persistent. Pileus yellowish with age; pileus centre cream; stipe white; basidia with 4 sterigmata; spores globose to sub-globose, 9–11 × 9–11 µm, Q = 1.0–1.13, Qm = 1.06 ± 0.03 A. alboumbelliformis Striation on pileus 0.1–0.3 R; annulus fragile 4 4. Pileus centre cream to alutaceous cream, striation 0.25 R. Annulus white; stipe squamules white in snakeskin pattern, basidia mainly with 2 sterigmata; spores globose to subglobose, 8.5–10.5 × 8.5–9.5 µm, Q = 1.04–1.14, Qm = 1.07 ± 0.06 A. chepangianoides Pileus centre white, sometimes with yellowish, yellowish or brownish tint, striation 0.1–0.15R. Annulus with age yellowish; stipe squamules white, bruising yellowish to greyish; basidia with 4 sterigmata; spores sub- globose to broadly ellipsoid, 9.5–12.5 × 8.5–11.5 µm, Q = 1.07–1.28, Qm = 1.17 ± 0.06 A . chepangiana 5. Striation on pileus 0.3–0.6 R; spores globose to subglobose 6 Striation on pileus less than 0.3 R; spores subglobose to broadly ellipsoid 7 6. Pileus to 140 mm wide, with broad umbo, striation 0.3–0.4 R, centre first greyish-orange soon golden brown. Stipe white, smooth to slightly fibrous; volval surface not cracking into brownish patches; annulus membranous, basidiospores 9–12 × 8.5–11.5 µm, Qm: 1.04 ± 0.06 A. fulvocaesarea Pileus 60–100 mm wide, slightly depressed, non-umbonate, striation 0.5–0.6 R, centre reddish-brown. Stipe pale cream, ± darker squamules; anulus fragile; basidiospores 8.5–11 × 8.5–10.5 µm, Qm: 1.06 ± 0.04 A. aporema 7. Pileus to 150 mm in diameter, centre yellowish cream, olive-grey to yellowish brown; stipe squamules white to dirty white; volval surface not cracking into brownish patches; basidiospores 9.5–14.5 × 8–12.5 µm, Qm = 1.12 ± 0.08 A. pseudoprinceps Pileus to 200 mm in diameter, centre brown to yellowish-brown; stipe squamules white, volva often cracking and peeling into thin patches; basidiospores 8–11 × 7.5–10 µm, Qm = 1.08 ± 0.05 A. princeps 8. Lamellar edge concolorous with lamellae 9 Lamellar edge yellowish or reddish orange 11 9. Pileus uniformly yellowish. Pileus 40–100 mm in diameter, striation 0.3–0.5 R; lamellae yellowish; stipe yellowish to yellow, squamules yellow to orange buff; annulus yellow; basidiospores broadly ellipsoid to ellipsoid, 8.5–13.5 × 6–8.5 µm, Qm = 1.41 ± 0.6 A. kitamagotake Pileus centre and margin of different colours 10 10. Pileus centre red to orange red to yellow, margin yellow. Pileus 60–150 mm in diameter, striation 0.3–0.5R; lamellae cream to yellowish; stipe yellowish, stipe squamules yellow to orange-yellow in irregular snakeskin pattern; annulus yellow; basidiospores ellipsoid, 7.5–11 × 5.5–7.5 µm, Qm = 1.49 ± 0.13 A. hemibapha Pileus centre yellowish to yellowish orange. Pileus 60–100 mm in diameter, margin pale yellow to maize yellow, striation 0.3–0.4R; stipe yellowish to cream, stipe squamules below annulus yellow; lamellae whitish, edge colour unknown probably concolorous; annulus yellowish white to whitish; basidiospores broadly ellipsoid to ellipsoid, 9–12 × 6–8 µm, Qm 1.39 ± 0.13 A. malayensis 11. Lamellar edge reddish orange. Pileus center orange red to brownish-red to red, margin orange to deep yellow, 60–90(130) mm in diameter; striation 0.4–0.5 R; stipe yellowish, stipe squamules orange to orange-red, often in a snakeskin pattern; lamellae yellow, edge light orange to reddish-orange; annulus, dark orange to reddish-orange; basidiospores 6.5–9 × 5.5–7 µm, Qm 1.22 ± 0.07. A. rubromarginata Lamellar edge in yellow colours 12 12. Pileus center brownish-orange to reddish-orange; margin orange yellow, 60–110 mm in diameter; striation 0.2–0.35 R. Lower pileipellis with yellowish zone (0.7–0.9 mm); stipe yellowish white, stipe squamules reddish-orange to dark orange; lamellae yellowish-white to light yellow, annulus yellowish-white to orange; basidiospores 7.5–10 × 6.5–8 µm, Qm: 1.23 ± 0.08 A. subhemibapba Pileus center orange-yellow to ochre yellow, margin paler or buff, 60–90 mm in diameter; striation 0.4–0.5 R. Lamellae whitish, stipe yellowish, stipe squamules orange-buff; annulus orange buff, membranous; basidiospores 7.5–9 × 5.8–7 µm, Qm: 1.26 ± 0.06 A. javanica Declarations Acknowledgements Urmas Kõljalg, Irja Saar and staff at Institute of Ecology and Earth Sciences, Department of Plant Pathology, University of Tartu, Estonia, are gratefully thanked for carrying out DNA analyses and uploading to the PlutoF platform. The encouragements from and many discussions with Andy Taylor of the James Hutton Institute are sincerely appreciated. Finally, Yang-Yang Cui and Zhu Liang Yang are heartfully thanked for their constructive suggestions and for allowing us to use unpublished data. Field work in Laos was partly GEF-financed under the UNDP-FAO Agrobiodiversity Project (UNTS/LAO/015/GEF), partly by the Darwin Initiative Project “Building mycological capacity for sustainable resources management in Lao PDR” of The James Hutton Institute and partly through private means. Drafts of the manuscript were written by OSP with regular comments and suggestions from TL. Both authors have read and approved the final manuscript. Datasets generated and analyzed during the current study are available from the corresponding author on reasonable request. The authors declare no competing interests related to this study. 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Bong, Mixay, Yai, Lethong, Gnordphe and Poua-Xai villages, Phoukhout District, Xieng Khouang Province, Lao PDR. https://data.opendevelopmentmekong.net/dataset/a136ed27-71c2-4e9b-b3fe-767c80d62c69/resource/ac76e8e4-48e3-45d0-8018-6c1ae52f4ae9/download/wildmusroomsurveyreport2014final.pdf Qin J, Yang ZL 2016 Cyptotrama ( Physalacriaceae , Agaricales ) from Asia Fungal Bio120(4):513–529. doi: 10.1016/j.funbio.2016.01.00 Reid DA (1978) New species of Amanita ( fungi ) from Australia. The Victorian Naturalist. 95:47–49. https://www.biodiversitylibrary.org/item/123295#page/54/mode/1up Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol., 61:539–542. https://doi.org/10.1093/sysbio/sys029 Sysouphanthong P, Bouamanivong S, Salichanh T, Xaybouangeun N, Sucharitakul P, Osathanunkul M, Suwannapoom C (2018) Leucoagaricus houaynhangensis ( Agaricaceae ), a new yellowish-green species from Lao People’s Democratic Republic. Chiang Mai Journal of Science 45 (3): 1287–1295 Sysouphanthong P, Bouamanivong, S Salichang T (2017) Some mushrooms found in Huayyang Preserves. Biotechnology and Ecology Institute, Ministry of Science and Technology, Vientiane Sysouphanthong P, Thongklang N (2022) Two new species of Leucoagaricus ( Agaricaceae ) from Lao People’s Democratic Republic. Current Research in Environmental & Applied Mycology (Journal of Fungal Biology) 12(1): 65–74. doi 10.5943/cream/12/1/6 Sysouphanthong P, Thongklang N, Liu Y, Vellinga E. (2022) Three new species of Cystolepiota from Laos and Thailand. Diversity14(6):449. https://doi.org/10.3390/d14060449 Takahashi H (2004) Two new species of Agaricales from southwestern islands of Japan, Mycoscience 45(6):372 Tamura K, Stecher G, Kumar S. MEGA11 (2021) Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution. 38(7): 3022–27. https://doi.org/10.1093/molbev/msab120 Tang L-P, Lee S-S, Zeng N-K, Cai Q, Zhang P, Zhu L, Yang ZL (2017) Notes on Amanita section Caesareae from Malaysia. Mycologia 109(4):557–567. https://doi:10.1080/00275514.2017.1394789 Terashima Y, Takahashi H, Taneyama Y (eds.) (2016) The fungal flora in southwestern Japan: Agarics and boletes [English and Japanese text]. Tokai University Press Thongbai B, Tulloss RE, Miller SL, Hyde KD, Chen J, Zhao R, Raspé O (2016) C, Phytotaxa 286(4):211–231. http://dx.doi.org/10.11646/phytotaxa.286.4.1 211 Tulloss RE, Bhandary HR (1992) Amanita chepangiana – a new species from Nepal. Mycotaxon 43: 25–31. https://www.researchgate.net/publication/287997935 Tulloss RE, Rodríguez-Caycedo C (2011) Amanita workshop, NJMA. DOI: 10.13140/RG.2.1.1138.4406 Tulloss RE, Yang ZL. (eds) 2024) [multiple text] Amanitaceae studies. http://www.amanitaceae.org. Accessed 30 January 2024 Tulloss RE, Yang ZL (2009) Notes on Amanita section Caesareae, Torrendia, and Amarrendia ( Agaricales , Amanitaceae ) with provisional division into stirpes and annotated world key to species of the section, Working Paper. DOI: 10.13140/RG.2.1.1504.7767. https://www.researchgate.net/publication/254084706 van de Peppel LJJ, Aime MC, Læssøe T, Pedersen OS, Coimbra VRM, Kuyper TW, Stubbe D, Aanen D, Baroni TJ (2022) Four new genera and six new species of lyophylloid agarics ( Agaricales , Basidiomycota ) from three different continents. Mycol Progress 21, 85. https://doi.org/10.1007/s11557-022-01836-7 Vidal J (1958) Noms vernaculaires de plantes (lao, mèo, kha) en usage au Laos. Bulletin de l'École française d'Extrême-Orient, Paris 49–2:435–608. https://www.persee.fr/doc/befeo_0336-1519_1959_num_49_2_1493 Voitk A, Saar I, Lodge DJ, Boertmann D, Berch SM, Larsson E (2020) New species and reports of Cuphophyllus from northern North America compared with related Eurasian species, Mycologia, 112:2, 438–452. https://DOI: 10.1080/00275514.2019.1703476 Vrinda KB, Pradeep CK, Kumar SS (2005) Occurrence of a lesser known edible Amanita in the Western Ghats of Kerala. Mushroom Res. 14:5–8 Wang R, Herrera M, Xu W, Zhang P, Moreno JP, Colinas C, Yu F (2022) Ethnomycological study on wild mushrooms in Pu’er Prefecture, Southwest Yunnan, China. J Ethnobiol Ethnomed 18:55. https://doi.org/10.1186/s13002-022-00551-7 Yang ZL (2015) Atlas of the Chinese species of Amanitaceae [In Chinese]. ISBN 9787030436276, Science Press, Beijing Yang ZL, Cai Q, Cui, YY (2018) Phylogeny, diversity and morphological evolution of Amanitaceae . Biosyst Ecol Ser 34:359–380 Supplementary Files Supplementarytable1AmanitaITSsequencedata.docx Cite Share Download PDF Status: Published Journal Publication published 05 Oct, 2024 Read the published version in Mycological Progress → Version 1 posted Reviewers agreed at journal 18 May, 2024 Reviewers invited by journal 18 May, 2024 Editor invited by journal 15 May, 2024 Editor assigned by journal 13 May, 2024 First submitted to journal 10 May, 2024 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-4401254","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":303884436,"identity":"f6a99a20-d4a6-4fb5-bb04-6f3081fd6a2b","order_by":0,"name":"Ole Sparre Pedersen","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAxElEQVRIiWNgGAWjYDACdjDJLAciDzwgSgszhDQGa0kgRUtiA4giSgs/M/PhzwUV1unzww4/BNpiJ6fbQECLZDNbmvSMM+m5G2+nGQC1JBubHSCgxeAwjxkzb9vh3I2zE0BaDiRuI6yF//Nn3n+H0w1np38gVgsPgzRvw+EEeekcIm0B+sVMesaxdMMN0jkFBxIMiPALP3vz488FNdby8rPTN3/4UGEnR1ALCICjxgCs0oAI5XAt8g1Eqh4Fo2AUjIKRBwDR/0HpDaBbZgAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0009-0000-1271-4607","institution":"Retired","correspondingAuthor":true,"prefix":"","firstName":"Ole","middleName":"Sparre","lastName":"Pedersen","suffix":""},{"id":303884437,"identity":"e56f7417-f15c-495f-ae6c-2f2bf58cb122","order_by":1,"name":"Thomas Læssøe","email":"","orcid":"","institution":"University of Copenhagen Globe Institute: Kobenhavns Universitet Globe Institute","correspondingAuthor":false,"prefix":"","firstName":"Thomas","middleName":"","lastName":"Læssøe","suffix":""}],"badges":[],"createdAt":"2024-05-10 14:20:24","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4401254/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4401254/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11557-024-01999-5","type":"published","date":"2024-10-05T15:58:00+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":57359393,"identity":"41556a38-8ee8-4bf8-a282-64a1d78da717","added_by":"auto","created_at":"2024-05-29 14:57:34","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":781746,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree of ITS derived sequences from Bayesian Interference and Maximum Likelihood analyses of sect. \u003cem\u003eCaesareae\u003c/em\u003e. Bayesian posterior probabilities (first value) ≥ 0.95 and bootstrap support values for Maximum Likelihood (second value) ≥ 70 are placed near the branches. The ML tree is rooted to \u003cem\u003eA. pseudosychnopyramis \u003c/em\u003efrom sect.\u003cem\u003e Amanita\u003c/em\u003e. Newly generated Lao sequences used in this study are highlighted in blue. Type specimens are annotated with (T) and proposed new TYPE species are indicated in bold script.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/bf778b679b5be87b87e2ded3.png"},{"id":57360091,"identity":"ba32b50a-8f92-4db7-857e-837cd4d2faec","added_by":"auto","created_at":"2024-05-29 15:05:34","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1260790,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAmanita\u003c/em\u003e c\u003cem\u003ehepangianoides\u003c/em\u003e \u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003ee\u003c/strong\u003e\u003cem\u003e \u003c/em\u003eTYPE,\u003cem\u003e \u003c/em\u003eHNL501136: \u003cstrong\u003ea\u003c/strong\u003e Basidiomata in situ; \u003cstrong\u003eb\u003c/strong\u003e Lamellar trama: mediostratum and lateral stratum; \u003cstrong\u003ec\u003c/strong\u003e Inner volva; \u003cstrong\u003ed\u003c/strong\u003e Hymenium and subhymenium; \u003cstrong\u003ee\u003c/strong\u003e Basidiospores. \u003cstrong\u003ef\u003c/strong\u003e HNL501150 (paratype): basidiomata in situ. Scale bars: \u003cstrong\u003ea, f\u003c/strong\u003e = 30 mm; \u003cstrong\u003eb, \u003c/strong\u003e= 100 μm; \u003cstrong\u003ec\u003c/strong\u003e = 30 μm\u003cstrong\u003e d, e\u003c/strong\u003e = 10 μm\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/78cca8ea5817962dc9fb5cad.png"},{"id":57359395,"identity":"06fdd312-583f-4386-a655-49ba2281de71","added_by":"auto","created_at":"2024-05-29 14:57:34","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":917487,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAmanita\u003c/em\u003e \u003cem\u003efulvocaesarea \u003c/em\u003e\u003cstrong\u003ea-e \u003c/strong\u003eTYPE, HNL500511: \u003cstrong\u003ea\u003c/strong\u003e Basidiomata in situ; \u003cstrong\u003eb\u003c/strong\u003e Lamellar trama: mediostratum and lateral stratum; \u003cstrong\u003ec\u003c/strong\u003e Longitudinal section of inner part of volva; \u003cstrong\u003ed\u003c/strong\u003eHymenium and subhymenium;\u003cstrong\u003e e\u003c/strong\u003e Basidiospores. \u003cstrong\u003ef\u003c/strong\u003e HNL501130 (paratype): basidiomata in situ. Scale bars: \u003cstrong\u003ea, f\u003c/strong\u003e = 30 mm; \u003cstrong\u003eb\u003c/strong\u003e = 100 μm, \u003cstrong\u003ec\u003c/strong\u003e = 30 μm\u003cstrong\u003e d, e\u003c/strong\u003e = 10 μm\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/160613d176d56eff93f31f94.png"},{"id":57360094,"identity":"c6b89490-abcd-4e65-af74-af2b642244f3","added_by":"auto","created_at":"2024-05-29 15:05:34","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":380587,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAmanita\u003c/em\u003e \u003cem\u003epseudoprinceps \u003c/em\u003e\u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003ec\u003c/strong\u003e\u003cem\u003e \u003c/em\u003eHNL500333: \u003cstrong\u003ea\u003c/strong\u003e Basidioma in habitat; \u003cstrong\u003eb\u003c/strong\u003e Hymenium and subhymenium; \u003cstrong\u003ec\u003c/strong\u003e Basidiospores. \u003cstrong\u003ed\u003c/strong\u003e HNL500044: Basidiomata in situ. Scale bars: \u003cstrong\u003ea, d-e\u003c/strong\u003e = 30 mm; f= 10 mm; \u003cstrong\u003eb, c\u003c/strong\u003e = 10 µm\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/b412604c28c8cad59702bc82.png"},{"id":57360093,"identity":"1ad58c2f-1e62-4c7f-b0ff-4bbc187e645b","added_by":"auto","created_at":"2024-05-29 15:05:34","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":390288,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAmanita\u003c/em\u003e \u003cem\u003erubromarginata \u003c/em\u003e\u003cstrong\u003ea\u003c/strong\u003e–\u003cstrong\u003ed\u003c/strong\u003e\u003cem\u003e \u003c/em\u003eHNL500859: \u003cstrong\u003ea-b\u003c/strong\u003e Basidiomata in situ; \u003cstrong\u003ec\u003c/strong\u003e Basidium; \u003cstrong\u003ed\u003c/strong\u003e Basidiospores. \u003cstrong\u003ee\u003c/strong\u003eHNL502458: basidiomata in habitat. Scale bars: \u003cstrong\u003ea, b, e\u003c/strong\u003e = 30 mm; \u003cstrong\u003ec, d\u003c/strong\u003e= 10 µm\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/9d706c3a4eebae549ad36745.png"},{"id":57359398,"identity":"06b57492-f333-4dea-90b1-3fa6403f94d5","added_by":"auto","created_at":"2024-05-29 14:57:34","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":302797,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAmanita subhemibapha\u003c/em\u003e \u003cstrong\u003ea-d\u003c/strong\u003e HNL502059: \u003cstrong\u003ea-b\u003c/strong\u003e Basidiomata, \u003cstrong\u003ec\u003c/strong\u003e Hymenium and subhymenium, \u003cstrong\u003ed\u003c/strong\u003e Basidiospores, \u003cstrong\u003ee\u003c/strong\u003e HNL501890: basidiomata in situ. Scale bars: \u003cstrong\u003ea, e\u003c/strong\u003e = 30 mm; \u003cstrong\u003eb\u003c/strong\u003e= 10 mm\u003cstrong\u003e c, d\u003c/strong\u003e = 10 µm\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/23caa2024d2f0636dd176ffa.png"},{"id":66097413,"identity":"b0538eee-92a7-4764-b862-014ee2be072f","added_by":"auto","created_at":"2024-10-07 16:14:05","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":6580239,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/8ee3632d-94f7-49c6-9241-5b60fcc0bea9.pdf"},{"id":57359392,"identity":"05cdc37d-f823-467c-a313-735872aa001d","added_by":"auto","created_at":"2024-05-29 14:57:34","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":76173,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarytable1AmanitaITSsequencedata.docx","url":"https://assets-eu.researchsquare.com/files/rs-4401254/v1/2aed012224a0153b0db3f0fb.docx"}],"financialInterests":"","formattedTitle":"Amanita species within section Caesareae from the highlands of Laos","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe genus \u003cem\u003eAmanita\u003c/em\u003e Pers. (1797: 65) (\u003cem\u003eAmanitaceae, Agaricales)\u003c/em\u003e is widely occurring in temperate, subtropical, and tropical regions and contains approximately 1,200 binomials of which some 650 are currently accepted species (Yang et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Tulloss and Yang \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Mycorrhizal tree partners are foremost with \u003cem\u003eFagaceae\u003c/em\u003e, \u003cem\u003ePinaceae\u003c/em\u003e, \u003cem\u003eDipterocarpaceae\u003c/em\u003e and \u003cem\u003eBetulaceae\u003c/em\u003e (Tulloss and Yang \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), all with members in Laos (Newman et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Currently, three subgenera are accepted within \u003cem\u003eAmanita\u003c/em\u003e - \u003cem\u003eLepidella, Amanitina\u003c/em\u003e and \u003cem\u003eAmanita\u003c/em\u003e - with subgenus \u003cem\u003eAmanita\u003c/em\u003e having four sections: \u003cem\u003eAmanita, Amarrendiae, Caesareae\u003c/em\u003e, and \u003cem\u003eVaginatae\u003c/em\u003e (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This study is limited to identified Lao specimens within sect. \u003cem\u003eCaesareae\u003c/em\u003e. This section, when agaricoid, is characterized by fruitbodies with a striate-sulcate pileus margin, truncate lamellulae, annulate stipes, a non-bulbous stipe base with a saccate volva, inamyloid basidiospores and presence of clamps (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The Amanitaceae.org website lists 101 binomials within sect. \u003cem\u003eCaesareae\u003c/em\u003e of which 71 have a detailed description (Tulloss and Yang \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Sect. \u003cem\u003eCaesareae\u003c/em\u003e has an almost cosmopolitan distribution with many species found in South and East Asia and in recent years several new species have been published from this region foremost from China, where for example six new species were added by Cui et al. (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLaos lies within the Indo-Burma Biodiversity Hotspot (Myers \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1988\u003c/span\u003e) and boasts a relative high forest coverage of 56%, spanning from dipterocarp dominated forests at lower altitudes to broadleaved-coniferous and evergreen forests at higher altitudes (MAF 2021). These forest types accommodate numerous ectomycorrhizal fungi, including many \u003cem\u003eAmanita\u003c/em\u003e species (L\u0026aelig;ss\u0026oslash;e et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePatouillard (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1928\u003c/span\u003e) was probably the first to publish on Laotian macrofungi among mainly aphyllophoroid species from Vietnam. Another early publication mentioning macrofungi in Laos was the Flora of Laos by Vidal (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1958\u003c/span\u003e), a French botanist, who included 33 macrofungal species. Much later, this was followed-up by a non-timber forest products handbook (NAFRI 2007), which included 29 common, edible and medicinal fungal species. Recent surveys by mycologists have been carried out at biodiversity conservation areas (Sysouphanthong et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Lee et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hong et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Lee et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), at local community forests (Pedersen and Thammavong \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and at local markets (Łuczaj et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Macrofungal species described based on material from Laos are rather few, but increased in recent years and includes \u003cem\u003ePhellinus stratosus\u003c/em\u003e Pat. (Patouillard \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1928\u003c/span\u003e), \u003cem\u003eCyptotrama angustispora\u003c/em\u003e Zhu L. Yang \u0026amp; J. Qin and \u003cem\u003eC. glabra\u003c/em\u003e Zhu L. Yang \u0026amp; J. Qin (Qin and Yang \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), \u003cem\u003eLeucoagaricus houaynhangensis\u003c/em\u003e Sysouphanthong (Sysouphanthong et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), \u003cem\u003eGanoderma nasalaense\u003c/em\u003e Hapuar., Pheng. et K.D. Hyde (Hapuarachchi et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), \u003cem\u003eHebeloma parvisporum\u003c/em\u003e O. S. Pedersen, L\u0026aelig;ss\u0026oslash;e, Beker \u0026amp; Eberh. (Eberhardt et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), \u003cem\u003eCystolepiota pyramidalis\u003c/em\u003e Sysouph. \u0026amp; Thongkl. and \u003cem\u003eC. rhodella\u003c/em\u003e Sysouph. \u0026amp; Thongkl. (Sysouphanthong et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), \u003cem\u003eNigrocarnea radicata\u003c/em\u003e O. S. Pedersen \u0026amp; L\u0026aelig;ss\u0026oslash;e (van de Peppel et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), \u003cem\u003eLeucoagaricus flavus\u003c/em\u003e Sysouph. \u0026amp; Thongkl. and \u003cem\u003eL. griseosquamosus\u003c/em\u003e Sysouph. \u0026amp; Thongkl. (Sysouphanthong and Thongklang \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and \u003cem\u003eL. laosensis\u003c/em\u003e Sysouph. (Manawasinghe et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). L\u0026aelig;ss\u0026oslash;e et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) published the first comprehensive introductory field guide on edible, poisonous and medicinal mushrooms of Northern Laos including 24 \u003cem\u003eAmanita\u003c/em\u003e species of which three were from sect. \u003cem\u003eCaesareae\u003c/em\u003e. The authors noted that many specimens need supporting molecular analyses to confirm their identity or make the description of new species possible. This study is the first of its kind from Laos, with the aim to document Lao species of genus \u003cem\u003eAmanita\u003c/em\u003e sect. \u003cem\u003eCaesareae\u003c/em\u003e, using morphological characters, molecular phylogenetic evidence and ecological data.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003eBasidiomata were collected during the monsoon season from May to September from rich mixed forests, mainly dominated by \u003cem\u003eFagaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e in the highland areas of Xieng Khouang and Vientiane provinces, Laos (Lao People's Democratic Republic). Specimens were deposited in the National Herbarium of Laos (HNL) with duplicates of some collections in C. Index Fungorum (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.IndexFungorum.org\u003c/span\u003e\u003cspan address=\"http://www.IndexFungorum.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), the International Index of Plant Names (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ipni.org\u003c/span\u003e\u003cspan address=\"https://www.ipni.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and Amanitaceae.org (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.Amanitaceae.org\u003c/span\u003e\u003cspan address=\"http://www.Amanitaceae.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) were used as sources for taxonomic names and nomenclature.\u003c/p\u003e \u003cp\u003eMacroscopic characters were described from fresh material with field notes and photographs. Pileal sulcation is annotated as the ratio (R) of the length of the striations from the pileal margin to the centre of the pileus. Stipe length is from the volval base. Colour codes follow Kornerup and Wanscher (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1981\u003c/span\u003e). Microscopic structures were observed with light microscopy (Brunel SP500 with OMAX 14MP mount microscope camera) from the dried specimens after sectioning and mounting in H\u003csub\u003e2\u003c/sub\u003eO and 3% KOH solution and where necessary stained with Congo Red. Basidiospores were tested for amyloidity with Melzer\u0026rsquo;s reagent and were measured in 3% KOH. The abbreviation (n/m/p) means n basidiospores measured from m basidiomata from p collections, using a range notation of the form (a\u0026ndash;) b\u0026ndash;c (\u0026ndash;d), the range b\u0026ndash;c contains a minimum of 90% of the measured values, extreme values (a, d) are given in parentheses, m and av. indicate average values, \u0026plusmn; standard deviation; Q presents the length/width ratio of spores in side view. Description terminology follows Bas (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1969\u003c/span\u003e), Tulloss and Rodr\u0026iacute;guez-Caycedo (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and Tulloss and Yang (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Sizes of basidiomata are described as small, medium or large with pileus diameter of 30\u0026ndash;50 mm, 50\u0026ndash;90 mm, and 90\u0026ndash;150 mm, respectively (Bas \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1969\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDNA extraction and sequencing of the ITS1, 5.8S, ITS2 regions were carried out at Tartu University, Estonia and uploaded into PlutoF cloud database (Abarenkov et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), following the procedures referred to and described in Voitk et al. (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Lao sequences were blasted in Unite website, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://unite.ut.ee/analysis.php\u003c/span\u003e\u003cspan address=\"https://unite.ut.ee/analysis.php\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (K\u0026otilde;ljalg et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e and Abarenkov et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and at NCBI website, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.ncbi.nlm.nih.gov/blast\u003c/span\u003e\u003cspan address=\"http://www.ncbi.nlm.nih.gov/blast\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e and related sequences (supplementary table 1) were retrieved for phylogenetic analyses. Priority was given to similar species, as well as to species representing neighbouring clades within sect. \u003cem\u003eCaesareae\u003c/em\u003e described in recent publications (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Kumla et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The final ingroup dataset consisted of 38 sequences of which 12 new ITS sequences were from Laos, and \u003cem\u003eA. pseudosychnopyramis\u003c/em\u003e from sect. \u003cem\u003eAmanita\u003c/em\u003e was selected as outgroup. Supplied by the NGPhylogeny website \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.phylogeny.fr/index.cgi\u003c/span\u003e\u003cspan address=\"http://www.phylogeny.fr/index.cgi\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (Lemoine et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) sequences was aligned with MUSCLE (Edgar \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and MAFFT v.7.4 (Katoh and Standley \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and inspected for reverse sequences in BioEdit (Hall \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e); poorly aligned positions were excluded applying the BMGE (Block Mapping and Gathering with Entropy) software (Criscuolo and Gribaldo, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), Maximum Likelihood (ML) phylogenetic analysis were performed by PhyML v.3.3 (Guindon et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2010\u003c/span\u003e)\u0026thinsp;+\u0026thinsp;Smart Model Selection (SMS) (Lefort et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and by Bayesian Interference (BI) analysis by MrBayes v. 3.2.7 (Ronquist et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The final trees were edited and visualized using Interactive Tree Of Life (iTOL) v.6 (Letunic and Bork \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and Microsoft Office. The BMGE selected 532 and removed 998 characters. MrBayes for BI was performed with 1.000.000 generations and otherwise default settings. The ML tree was examined by HKY85 substitution model with 1,000 replicates, otherwise default settings resulting in the log-likelihood tree value of -3698.83699 for the best scoring tree. The phylogram topology derived from BI and PhyML were similar and only the ML tree is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. In comparison, while running ML in MEGA11 (Tamura et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) a similar topology and bootstrap values were observed. A branch was considered strongly supported if it had ML bootstrap support\u0026thinsp;\u0026ge;\u0026thinsp;85% and posterior probability (PP)\u0026thinsp;\u0026ge;\u0026thinsp;0.95. Dataset and final alignments for \u003cem\u003eA. fulvocaesarea\u003c/em\u003e and \u003cem\u003eA. chepangianoides\u003c/em\u003e have been deposited in TreeBase (No. xxxxxxx/xxxxxxx and xxxxxxxxxxxx/xxxxxxx, respectively.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eTaxonomy\u003c/h2\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eEtymology\u003c/strong\u003e \u003cp\u003eEpithet refers to the similarity to \u003cem\u003eAmanita chepangiana\u003c/em\u003e Tulloss \u0026amp; Bhandary.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eHolotype\u003c/b\u003e: Laos. Xieng Khouang province, Phoukhout district, Ngodphae village, Na Phouak hamlet, 19\u0026deg;45'43.8\"N, 103\u0026deg;15'22.5\"E, 1385 m asl., in broadleaved forest with \u003cem\u003eQuercus\u003c/em\u003e spp. \u003cem\u003eCastanea\u003c/em\u003e spp. and \u003cem\u003eLithocarpus\u003c/em\u003e spp., 23 May 2016, O. Sparre Pedersen and T. L\u0026aelig;ss\u0026oslash;e, OSP20160523\u0026ndash;013, HNL501136, as \u003cem\u003eA.\u003c/em\u003e? \u003cem\u003echepangiana\u003c/em\u003e in L\u0026aelig;ss\u0026oslash;e et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), isotype C-F-169624 (C), GenBank: ITS\u0026thinsp;=\u0026thinsp;xxxxxxxxx\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eDiagnosis\u003c/strong\u003e \u003cp\u003e \u003cem\u003eAmanita chepangianoides\u003c/em\u003e is characterized by white basidiomata except the alutaceous-cream centre of the pileus, a relative short pileal margin striation (0.25 R), a snake-skin pattern on the stipe and globose to subglobose spores (8.5\u0026ndash;10.5 \u0026times; 8.5\u0026ndash;9.5 \u0026micro;m) and presence of clamps.\u003c/p\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eDescription\u003c/h2\u003e \u003cp\u003e \u003cem\u003eBasidiomata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea, f) medium to large. \u003cem\u003ePileus\u003c/em\u003e 90\u0026ndash;130 mm in diam, initially almost spheric then convex and centrally depressed, slightly viscid, cream (4A2\u0026ndash;3) to alutaceous cream (5B4\u0026ndash;7B4) at centre and whitish (5A1\u0026ndash;2) towards margin; velar remnants absent; margin striate (0.25 R), non-appendiculate; trama white (1A1), unchanging. \u003cem\u003eLamellae\u003c/em\u003e free or almost so, plentiful (approximately 100 reaching stripe), white (1A1), edge fimbriate, concolorous when fresh and when dried; \u003cem\u003elamellulae\u003c/em\u003e truncate, few of 2\u0026ndash;3 lengths. \u003cem\u003eStipe\u003c/em\u003e 160\u0026ndash;200 \u0026times; 10\u0026ndash;18 mm, white (1A1), cylindrical, slightly tapering upwards, basal bulb absent, covered by white (1A1) squamules, mostly in a snake-skin pattern; context white (1A1), unchanging; \u003cem\u003evolva\u003c/em\u003e large, ca. 45 mm high and 35 mm wide, flaring, saccate, white (1A1). \u003cem\u003eAnnulus\u003c/em\u003e apical, membranous, white (1A1), upper surface sometimes weakly striate, hanging, fugacious. \u003cem\u003eTaste\u003c/em\u003e not recorded. \u003cem\u003eSmell\u003c/em\u003e pleasant, not prominent. \u003cem\u003eSpore deposit\u003c/em\u003e not observed.\u003c/p\u003e \u003cp\u003e \u003cem\u003eLamellar trama\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). Mediostratum 20\u0026ndash;40 \u0026micro;m wide, composed of fairly abundant ellipsoid to fusiform to cylindrical, thin-walled inflated cells, 25\u0026ndash;72 \u0026times; 17\u0026ndash;32 \u0026micro;m; filamentous hyphae abundant, 1\u0026ndash;2 \u0026micro;m wide; lateral stratum 37\u0026ndash;55 \u0026micro;m wide, composed of ellipsoid to clavate inflated cells (15\u0026ndash;40 \u0026times; 10\u0026ndash;15 \u0026micro;m), filamentous hyphae scarce, 1\u0026ndash;2 my wide. \u003cem\u003eSubhymenium\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed) 25\u0026ndash;42 \u0026micro;m thick composed of 2\u0026ndash;3 layers of subglobose, ovoid, ellipsoid to clavate cells, 12\u0026ndash;20 \u0026times; 9\u0026ndash;14 \u0026micro;m. \u003cem\u003eLamellar edge\u003c/em\u003e sterile, composed of irregular inflated cells, subglobose to irregular, hyaline, thin-walled, 16\u0026ndash;29 \u0026times; 15\u0026ndash;25 \u0026micro;m; filamentous hyphae scattered, 0.5\u0026ndash;2 \u0026micro;m wide. \u003cem\u003eBasidia\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed) 22\u0026ndash;35 \u0026times; 8\u0026ndash;14 \u0026micro;m, clavate, mainly 2-spored, few 4-spored; sterigmata 3\u0026ndash;6 \u0026micro;m long. \u003cem\u003eBasidiospores\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ee) [40/1/1] (8\u0026ndash;) 8.5\u0026ndash;10.5 (\u0026ndash;11) \u0026times; (7.5\u0026ndash;) 8.5\u0026ndash;9.5 (\u0026ndash;10.5) \u0026micro;m (av. 9.6 \u0026times; 8.8 \u0026micro;m), Q\u0026thinsp;=\u0026thinsp;1.04\u0026ndash;1.14 (Qm\u0026thinsp;=\u0026thinsp;1.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06), globose to subglobose (few broadly ellipsoid), inamyloid, hyaline, thin-walled, smooth; apiculus 1\u0026ndash;2 \u0026micro;m my high and 1\u0026ndash;1.5 \u0026micro;m wide. \u003cem\u003ePileipellis\u003c/em\u003e 80\u0026ndash;120 \u0026micro;m thick; upper layer 25\u0026ndash;70 \u0026micro;m, gelatinized, composed of radically arranged, filamentous hyphae 2\u0026ndash;10 \u0026micro;m wide; lower layer 25\u0026ndash;50 \u0026micro;m composed of radically arranged filamentous hyphae, 4\u0026ndash;8 \u0026micro;m wide, hyaline to yellowish. \u003cem\u003eInterior of volva\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec) composed of longitudinally arranged elements; filamentous hyphae very abundant 1\u0026ndash;7 \u0026micro;m wide, few branching; inflated cells scattered to locally abundant, subglobose, broadly ellipsoid, ellipsoid to cylindric 25\u0026ndash;62 \u0026times; 15\u0026ndash;45 \u0026micro;m, hyaline to yellowish, thin-walled; gelatinized hyphae locally abundant. \u003cem\u003eStipe trama\u003c/em\u003e composed of longitudinally arranged, mainly cylindrical terminal cells, thin-walled, 30\u0026ndash;240 \u0026times; 14\u0026ndash;65 \u0026micro;m; filamentous hyphae scattered, 3\u0026ndash;10 \u0026micro;m wide. \u003cem\u003eAnnulus\u003c/em\u003e composed of interwoven filamentous hyphae and inflated cells; inflated cells rare, subglobose to ellipsoid, 25\u0026ndash;55 \u0026times; 24\u0026ndash;35 \u0026micro;m; filamentous hyphae abundant, 2\u0026ndash;8 \u0026micro;m wide. \u003cem\u003eClamps\u003c/em\u003e presumably present in all parts, but difficult to observe, seen with certainty on stem hyphae.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabitat\u003c/strong\u003e \u003cp\u003eScattered on rich soil in broadleaved highland forests under \u003cem\u003eFagaceae\u003c/em\u003e.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eAdditional specimen examined\u003c/strong\u003e \u003cp\u003eLaos. Xiang Khouang province, Phoukhout district, Ngodphae village, 19\u0026deg;45'43.3\"N, 103\u0026deg;15'23.0\"E, 1380 m asl., in broadleaved forest with \u003cem\u003eQuercus\u003c/em\u003e spp., \u003cem\u003eCastanea\u003c/em\u003e spp. and \u003cem\u003eLithocarpus\u003c/em\u003e spp., 23 May 2016, O. Sparre Pedersen and T. L\u0026aelig;ss\u0026oslash;e, OSP20160523-012 (HNL501150).\u003c/p\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDiscussion\u003c/h3\u003e\n\u003cp\u003eOur ITS phylogenetic analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) places \u003cem\u003eA. chepangianoides\u003c/em\u003e in a larger clade containing \u003cem\u003eA. chepangiana\u003c/em\u003e Tulloss \u0026amp; Bhandary, \u003cem\u003eA. fulvocaesarea\u003c/em\u003e O.S. Pedersen \u0026amp; T. L\u0026aelig;ss\u0026oslash;e (described as new in this study), \u003cem\u003eA. princeps\u003c/em\u003e Corner and Bas, \u003cem\u003eA. aporema\u003c/em\u003e Boedijn, \u003cem\u003eA. egregia\u003c/em\u003e D.A. Reid and \u003cem\u003eA. pseudoprinceps\u003c/em\u003e Y.-Y. Cui, Q. Cai \u0026amp; Zhu L. Yang.\u003c/p\u003e \u003cp\u003eMorphologically, the similar \u003cem\u003eA. chepangiana\u003c/em\u003e differs by sometimes having yellowish to greyish tinges at centre of the pileus and by subglobose to ellipsoid basidiospores (9.5\u0026ndash;12.5 \u0026times; 8.5\u0026ndash;11.5 \u0026micro;m, Q\u0026thinsp;=\u0026thinsp;1.07\u0026ndash;1.28, Qm\u0026thinsp;=\u0026thinsp;1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06) (Tulloss and Bhandary \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). \u003cem\u003eAmanita alboumbelliformis\u003c/em\u003e Y.-Y. Cui, Q. Cai \u0026amp; Zhu L. Yang is also similar by sometimes having a cream pileus centre, but it differs by having a longer pilea striation (0.3\u0026ndash;0.4 R), larger spores (9\u0026ndash;11 \u0026times; 9\u0026ndash;11 \u0026micro;m, Q\u0026thinsp;=\u0026thinsp;1.0\u0026ndash;1.13, Qm\u0026thinsp;=\u0026thinsp;1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03), yellowish tinges on the aging annulus and a greyish to brownish volva (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). \u003cem\u003eAmanita egregia\u003c/em\u003e has white basidiomata, but differs by having larger pilei and stipes (up to 200 mm), broadly ellipsoid spores and association with \u003cem\u003eEucalyptus\u003c/em\u003e in Australia (Reid \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e1978\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eAmanita fulvocaesarea\u003c/b\u003e O. S. Pedersen \u0026amp; T. L\u0026aelig;ss\u0026oslash;e, sp. nov. (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e)\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eMycoBank\u003c/strong\u003e \u003cp\u003eMB xxxxx.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eHolotype\u003c/b\u003e: Laos. Xiang Khouang province, Phoukhout district, Ngodphae village, Na Phouang hamlet 19\u0026deg;45'11.9\"N, 103\u0026deg;15'29.1\"E, 1135 m asl., on soil with \u003cem\u003eQuercus\u003c/em\u003e spp., \u003cem\u003eCastanopsis\u003c/em\u003e spp. and \u003cem\u003eLithocarpus\u003c/em\u003e spp., 11 May 2015, O. Sparre Pedersen and T. L\u0026aelig;ss\u0026oslash;e, OSP20150511\u0026ndash;007, HNL500511, as \u003cem\u003eA. princeps\u003c/em\u003e in L\u0026aelig;ss\u0026oslash;e et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), isotype C-F-169623 (C), GenBank Acc. No.: ITS xxxxxxxxx.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eDiagnosis\u003c/b\u003e: \u003cem\u003eAmanita fulvocaesarea\u003c/em\u003e is characterized by the initial greyish-orange colours followed by golden brown hues, an umbonate pileus, distinct sulcation (0.3\u003cb\u003e\u0026ndash;\u003c/b\u003e0.4R), an apical annulus, white non-cracking volva, globose to subglobose basidiospores (9\u0026ndash;12 x 9\u0026ndash;11.5 \u0026micro;m) and presence of clamps.\u003c/p\u003e\n\u003ch3\u003eDescription\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eBasidiomata\u003c/em\u003e medium to large (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, f). \u003cem\u003ePileus\u003c/em\u003e up to 140 mm in diameter, first ovoid then convex to applanate with small broad umbo, dry to slightly viscid, centre first greyish-orange (5B4), with age golden brown (5D6) with tinges of greyish-ochraceous (5\u0026ndash;8B2) towards margin; veil remnants absent; margin sulcate (0.3\u0026ndash;0.4 R), non-appendiculate; trama white (1A1), unchanging. \u003cem\u003eLamellae\u003c/em\u003e free or almost so, whitish (1A1), when dry brownish-orange (6C5), appr. 100 reaching the stipe; \u003cem\u003elamellulae\u003c/em\u003e few of 2\u0026ndash;3 lengths, truncate; edge fimbriate, sterile, concolorous, when dried whitish-cream (4A1\u0026ndash;2). \u003cem\u003eStipe\u003c/em\u003e 150\u0026ndash;200 \u0026times;12\u0026ndash;20 mm, white (1A1) smooth to slightly fibrous, tapering towards apex, sometimes almost cylindrical; context white (1A1), hollow; basal bulb absent; \u003cem\u003evolva\u003c/em\u003e 30\u0026ndash;60 mm high and 20\u0026ndash;40 mm wide, saccate, thick, white (1A1), not cracking, flaring. \u003cem\u003eAnnulus\u003c/em\u003e white (1A1) on both sides, apical, big, pendent, but rather fugacious; upper surface non-striate. \u003cem\u003eTaste\u003c/em\u003e not recorded. \u003cem\u003eOdour\u003c/em\u003e indistinctive. \u003cem\u003eSpore deposit\u003c/em\u003e not observed.\u003c/p\u003e \u003cp\u003e \u003cem\u003eLamellar trama\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb) bilateral. \u003cem\u003eMediostratum\u003c/em\u003e 42\u0026ndash;63 \u0026micro;m wide, interwoven, inflated cells thin-walled, hyaline, scattered, fusiform, ellipsoid to clavate 18\u0026ndash;180 \u0026times; 7\u0026ndash;27 \u0026micro;m; filamentous hyphae abundant, 1\u0026ndash;5 \u0026micro;m wide; lateral stratum 36\u0026ndash;64 \u0026micro;m wide with abundant clavate to cylindrical, inflated cells 23\u0026ndash;38 \u0026times; 5\u0026ndash;11 \u0026micro;m; filamentous hyphae scarce, 1\u0026ndash;2 my wide. S\u003cem\u003eubhymenium\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed) 14\u0026ndash;34 \u0026micro;m thick, composed of 2\u0026ndash;3 layers of subglobose, ellipsoid to irregular cells, 8\u0026ndash;21 \u0026times; 6\u0026ndash;15 \u0026micro;m. \u003cem\u003eLamellar edge\u003c/em\u003e sterile composed of scattered subglobose, ellipsoid to cylindric, inflated, hyaline, thin-walled cells, 16\u0026ndash;36 \u0026times; 11\u0026ndash;18 \u0026micro;m and scattered filamentous, 1\u0026ndash;2.5 \u0026micro;m wide. \u003cem\u003eBasidia\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed) 42\u0026ndash;50 \u0026times; 15\u0026ndash;22 \u0026micro;m, clavate, 2- (few) and 4-spored, sterigmata 3\u0026ndash;6 \u0026micro;m long. \u003cem\u003eBasidiospores\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ee) [30/1/1] 9\u0026ndash;12 (\u0026ndash;12.5) \u0026times; 9\u0026ndash;11.5 (\u0026ndash;12) \u0026micro;m (av. 11.5 \u0026times; 11 \u0026micro;m), Q\u0026thinsp;=\u0026thinsp;1.01\u0026ndash;1.08, (Qm 1.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03); globose to subglobose, inamyloid, hyaline, thin-walled, smooth; apiculus 1.5\u0026ndash;2 \u0026micro;m high and 1\u0026ndash;2 \u0026micro;m wide. \u003cem\u003ePileipellis\u003c/em\u003e 135\u0026ndash;200 \u0026micro;m thick; upper layer 40\u0026ndash;140 \u0026micro;m, gelatinized, composed of radially arranged to interwoven hyphae, 4\u0026ndash;11 \u0026micro;m wide, hyaline to yellowish-brown; lower layer 30\u0026ndash;75 \u0026micro;m thick, composed of interwoven filamentous hyphae 1.5\u0026ndash;3.5 \u0026micro;m wide, hyaline to yellowish. \u003cem\u003eVolva\u003c/em\u003e dominantly composed of filamentous hyphae; interior surface (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec) composed of longitudinally orientated filamentous hyphae, 2\u0026ndash;4 \u0026micro;m wide, hyaline, thin-walled; outer surface of similar structure, but with gelatinized filamentous hyphae 2\u0026ndash;6 \u0026micro;m wide; inflated cells scattered, subglobose, fusiform to ellipsoid, 15\u0026ndash; 90\u0026times;10\u0026ndash;50 \u0026micro;m, hyaline, thin-walled. \u003cem\u003eStipe trama\u003c/em\u003e composed of longitudinally arranged, inflated, cylindrical to slightly clavate, hyaline, thin-walled terminal cells, 67\u0026ndash;215 \u0026times; 31\u0026ndash;45 \u0026micro;m; filamentous hyphae abundant, 3\u0026ndash;5 \u0026micro;m wide. \u003cem\u003eAnnulus\u003c/em\u003e composed of interwoven elements: outer part with inflated cells rare to scattered, subglobose to ellipsoid, 15\u0026ndash;34 \u0026times; 13\u0026ndash;23 \u0026micro;m, hyaline, thin-walled; filamentous hyphae very abundant 1\u0026ndash;5 \u0026micro;m wide, hyaline, thin-walled; inner part with radially arranged inflated cells, scattered to fairly abundant, hyaline, subglobose to elongate, 20\u0026ndash;44 \u0026times; 8\u0026ndash;24 \u0026micro;m; filamentous hyphae abundant, 1\u0026ndash;6 \u0026micro;m wide, hyaline, yellow to yellowish-brown, thin-walled. \u003cem\u003eClamps\u003c/em\u003e presumably present in all parts, but difficult to observe, seen with certainty on pileipellis hyphae and on stem hyphae.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabitat and distribution\u003c/strong\u003e \u003cp\u003eOn rich soil in broadleaved forests with members of \u003cem\u003eFagaceae\u003c/em\u003e (Laos) \u003cem\u003eFagaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e (China). Known from Laos (this study) and China (non-described).\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eAdditional specimen examined\u003c/strong\u003e \u003cp\u003eLaos. Xiang Khouang province, Phoukhout district, Ngodphae village, 19\u0026deg;45'43.3\"N, 103\u0026deg;15'23.0\"E, 1381 m asl., in broadleaved forest with \u003cem\u003eQuercus\u003c/em\u003e spp., \u003cem\u003eCastanopsis\u003c/em\u003e spp. and \u003cem\u003eLithocarpus\u003c/em\u003e spp., 23 May 2016, O. Sparre Pedersen and T. L\u0026aelig;ss\u0026oslash;e, OSP20160523\u0026ndash;009 (HNL501130). China. Yunan province, Puer, ca. 1400 m asl., in a forest with \u003cem\u003eLithocarpus\u003c/em\u003e and \u003cem\u003ePinus\u003c/em\u003e, 6 July 2015, Qing Cai 1349 (KUN-HKAS91972). The analyses of the Chinese taxa are based on Genbank ITS sequences, only.\u003c/p\u003e \u003c/p\u003e\n\u003ch3\u003eDiscussion \u003c/h3\u003e\n\u003cp\u003eOur phylogenetic analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) showed that three \u003cem\u003eAmanita\u003c/em\u003e specimens: HKAS91972 (ITS\u0026thinsp;=\u0026thinsp;MH508608), HKAS101390 (ITS\u0026thinsp;=\u0026thinsp;MH508603) and HKAS91096 (ITS\u0026thinsp;=\u0026thinsp;MH508607) from China, clustered with \u003cem\u003eA. fulvocaesarea\u003c/em\u003e with strong ML/PP support. These specimens were briefly mentioned by Cui et al. (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), but not assigned to any described taxon, due to insufficient morphological observations. For related species in the clade please see the commentary section of \u003cem\u003eA. chepangianoides\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eMorphologically \u003cem\u003eA. fulvocaesarea\u003c/em\u003e is similar to \u003cem\u003eA. pseudoprinceps and A. princeps\u003c/em\u003e. The former differs by a shorter pileal striation (0.2\u0026ndash;0.3 R) and subglobose to broadly ellipsoid spores (Q\u0026thinsp;=\u0026thinsp;1.02\u0026ndash;1.23, Qm\u0026thinsp;=\u0026thinsp;1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04) (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and in this study (Q\u0026thinsp;=\u0026thinsp;1.02\u0026ndash;1.25, Qm\u0026thinsp;=\u0026thinsp;1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08). The latter species differs by a larger (up to 200 mm) brownish to yellow pileus, shorter pileal striation (0.2\u0026ndash;0.25 R), a volval outer surface that often cracks and peels into thin brownish patches (Corner and Bas \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1962\u003c/span\u003e; Yang \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and the association with dipterocarps (Malaysia) (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). \u003cem\u003eAmanita aporema\u003c/em\u003e has some similarities, but it differs by relatively smaller basidiomata and darker coloured pilei (brown to reddish-brown, ca. 60\u0026ndash;100 mm in diam.) and longer striations, ca. 0.5 R (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eDescription\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eBasidiomata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, d-f) medium sized to large. \u003cem\u003ePileus\u003c/em\u003e 60\u0026ndash;140 mm in diam, initially hemispheric, then convex to applanate, slightly viscid, olive grey (1E2) to yellowish-brown (5D\u0026ndash;E8), darker towards centre and white to yellowish-cream (3\u0026ndash;4A3) to white (1A1) at margin; velar remnants absent; margin striate (0.2\u0026ndash;0.25 R), non-appendiculate; trama white (1A1), unchanging. \u003cem\u003eLamellae\u003c/em\u003e free or almost so, white (1A1) to whitish-cream (4A1\u0026ndash;2), plentiful (\u0026gt;\u0026thinsp;100 reaching stipe), up to 20 mm wide; \u003cem\u003elamellulae\u003c/em\u003e truncate, few of 5\u0026ndash;6 lengths. \u003cem\u003eStipe\u003c/em\u003e 70\u0026ndash;150 \u0026times; 10\u0026ndash;20 mm, cylindrical, basal bulb absent, white (1A1) with slightly darker fibrils; context white (1A1), unchanging, hollow; \u003cem\u003evolva\u003c/em\u003e large, saccate, 30\u0026ndash;60 mm high and 20\u0026ndash;40 mm wide, white (1A1), sometimes brownish (2B2-4) patches. \u003cem\u003eAnnulus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef) apical, skirt-like, membranous, 10\u0026ndash;20 mm high, white (1A1), partly striate on upper surface. \u003cem\u003eTaste\u003c/em\u003e not recorded. \u003cem\u003eSmell\u003c/em\u003e insignificant. S\u003cem\u003epore deposit\u003c/em\u003e not observed.\u003c/p\u003e \u003cp\u003e \u003cem\u003eBasidia\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eb) clavate, 28\u0026ndash;54 \u0026times; 10\u0026ndash;17 \u0026micro;m, 4-spored, sterigmata 1.5\u0026ndash;7 \u0026micro;m long. \u003cem\u003eBasidiospores\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ec) [120/3/3] (8.5\u0026ndash;) 9.5\u0026ndash;14.5 (\u0026ndash;18.5) \u0026times; (7.5\u0026ndash;) 8\u0026ndash;12.5 (\u0026ndash;13.5) \u0026micro;m (av. 12 \u0026times; 10.5 \u0026micro;m), Q\u0026thinsp;=\u0026thinsp;1.02\u0026ndash;1.25 (Qm\u0026thinsp;=\u0026thinsp;1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08), subglobose to broadly ellipsoid, very few globose, inamyloid, hyaline, thin-walled, smooth; apiculus 1.5\u0026ndash;2 \u0026micro;m high and 1\u0026ndash;1.5 \u0026micro;m wide. \u003cem\u003ePileipellis\u003c/em\u003e 95\u0026ndash;150 my thick; upper layer composed of radically arranged gelatinized filamentous hyphae, 1\u0026ndash;2 \u0026micro;m wide; lower layer composed of radically arranged, hyaline, filamentous hyphae, 4\u0026ndash;8 my wide. \u003cem\u003eInterior of volva\u003c/em\u003e composed of filamentous hyphae 1.5\u0026ndash;8 \u0026micro;m wide; inflated cells scattered, ellipsoid, hyaline, hyaline, thin-walled, scattered to locally abundant, 22\u0026ndash;55 \u0026times; 13\u0026ndash;25 \u0026micro;m. \u003cem\u003eStipe trama\u003c/em\u003e composed of longitudinally arranged, ellipsoid, clavate to cylindric, thin-walled terminal cells, 85\u0026ndash;170 \u0026times; 15\u0026ndash;60 \u0026micro;m; filamentous hyphae scattered to abundant, 2\u0026ndash;10 \u0026micro;m wide. \u003cem\u003eClamps\u003c/em\u003e not observed.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabitat and distribution\u003c/strong\u003e \u003cp\u003eScattered on soil in mixed forests with members of \u003cem\u003eFagaceae\u003c/em\u003e, \u003cem\u003eDipterocarpaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e or without the dipterocarp element, at altitudes 800\u0026ndash;1125 m, May to August. Known from southwestern China, Nepal, Laos and Thailand. Common at local markets in Laos.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecimens examined\u003c/strong\u003e \u003cp\u003eLaos. Vientiane province, Thoulakhom district, Vangheua village, 18\u0026deg;20'31.5\"N, 102\u0026deg;47'46.6\"E 790 m asl., in mixed forest with \u003cem\u003ePinus kesiya\u003c/em\u003e, \u003cem\u003eDipterocarpus\u003c/em\u003e spp. and \u003cem\u003eFagaceae\u003c/em\u003e, 19 May 2012, O. Sparre Pedersen, OSP20120519-001 (HNL500333). Ibid., 18\u0026deg;20'39.7\"N, 102\u0026deg;47'36.9\"E. 803 m asl., 6 July 2013, O. Sparre Pedersen, OSP20130706-004 (HNL500044). Xieng Khouang province, Pek district, Dong Village, 19\u0026deg;29'39.1\"N, 103\u0026deg;16'07.7\"E 1123 m asl., in mixed forest including \u003cem\u003ePinus kesiya\u003c/em\u003e, \u003cem\u003eKeteleeria everlyniana\u003c/em\u003e and \u003cem\u003eQuercus\u003c/em\u003e spp. 22 June 2014, O. Sparre Pedersen, OSP20140622\u0026ndash;007 (HNL500154).\u003c/p\u003e \u003c/p\u003e\n\u003ch3\u003eDiscussion \u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eAmanita pseudoprinceps\u003c/em\u003e was recently described from Yunnan, China, from a \u003cem\u003eFagaceae\u003c/em\u003e dominated mixed forest at 2300 m altitude (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The Laotian specimens (HNL500044, HNL500155 and HNL500333) match the type in terms of ecology, pileus and stipe colours, striation on pileus, apical placement of annulus, spore size as well as in our phylogenetic analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). A minor difference is that Laotian specimens have a hollow stipe in full length, compared to the holotype, which is only hollow in the centre. As discussed above, \u003cem\u003eA. pseudoprinceps\u003c/em\u003e can be easily confused with \u003cem\u003eA. princeps\u003c/em\u003e, which differs by a broader pileus (100\u0026ndash;200 mm), with light brown to yellowish-brown colours and with brownish cracking patches on the outer surface of the volva and association with dipterocarps (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). It also resembles \u003cem\u003eA. fulvocaesarea\u003c/em\u003e (described as new above), which is smaller, has a longer pileal striation (0.3\u0026ndash;0.4 R) and dominantly globose spores. The two other species shown in the clade (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), \u003cem\u003eA. aporema\u003c/em\u003e and \u003cem\u003eA. egregia\u003c/em\u003e differ from \u003cem\u003eA. pseudoprinceps\u003c/em\u003e by a brown to reddish-brown pileus and a white basidioma, respectively (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Reid \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e1978\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRecent work of Łuczaj et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) surveying markets in Luang Prabang, Laos, as well as an ethnobotanical survey in Yunnan, China by Wang et al. (\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) identified two taxa as \u003cem\u003eA. princeps\u003c/em\u003e (WA0000072256: ITS\u0026thinsp;=\u0026thinsp;MT 252585 and HKAS122502: ITS\u0026thinsp;=\u0026thinsp;ON394332, respectively). With strong bootstrap support in our study (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) these taxa should be referred to as \u003cem\u003eA. pseudoprinceps\u003c/em\u003e. See Cui et al. (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) for further misidentified specimens.\u003c/p\u003e \u003cp\u003e \u003cb\u003eAmanita rubromarginata\u003c/b\u003e Har. Takah., Mycoscience 45(6): 372 (2004) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e)\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eDescription\u003c/h2\u003e \u003cp\u003e \u003cem\u003eBasidiomata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea, b, f) medium sized. \u003cem\u003ePileus\u003c/em\u003e 45\u0026ndash;90 mm in diam, first almost cylindrical, then convex to applanate, centrally depressed, occasionally with broad umbo, dry or almost so, orange red (8A8) to brownish-red (9C\u0026ndash;D8) at centre and orange (5A6\u0026ndash;7), yellowish-orange (4AB7) to deep yellow (4A8) at margin; velar remnants absent; margin striate (0.4\u0026ndash;0.5 R), non-appendiculate, trama yellowish (4A4\u0026ndash;5). \u003cem\u003eLamellae\u003c/em\u003e free or almost so, plentiful (ca. 110 reaching stem), yellowish (2A7); edge fimbriate, light orange (5A4) to reddish-orange (7A7); \u003cem\u003elamellulae\u003c/em\u003e truncate, few of 3\u0026ndash;4 lengths. \u003cem\u003eStipe\u003c/em\u003e 70\u0026ndash;120 \u0026times; 7\u0026ndash;9 mm, cylindrical to slightly tapering upwards, basal bulb absent, hollow, yellow (2A7\u0026ndash;3A7) to light yellow (4A4\u0026ndash;5), partially covered by orange (5A6) to orange-red (8A8) squamules in a snake-skin pattern; context light yellow (4A4\u0026ndash;5); \u003cem\u003evolva\u003c/em\u003e thick, large, saccate, 35\u0026ndash;40 mm high and 25\u0026ndash;30 mm wide, whitish (2A2). \u003cem\u003eAnnulus\u003c/em\u003e subapical, membranous, hanging, dark orange (5A8) to reddish-orange (7A7), upper surface non-striate, edge dentate. \u003cem\u003eTaste\u003c/em\u003e not recorded. \u003cem\u003eSmell\u003c/em\u003e indistinct. \u003cem\u003eSpore deposit\u003c/em\u003e not observed.\u003c/p\u003e \u003cp\u003e \u003cem\u003eBasidia\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec) 9\u0026ndash;12 x 26\u0026ndash;37 \u0026micro;m clavate, 4-spored; sterigmata 2.5\u0026ndash;4.5 \u0026micro;m long; basal septa clamped. \u003cem\u003eBasidiospores\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ed) [40/2/2] 6.5\u0026ndash;9 (\u0026ndash;9.5) \u0026times; 5.5\u0026ndash;7 (\u0026ndash;7.5) \u0026micro;m (av. 7.5 \u0026times; 6.2 \u0026micro;m), Q: 1.13\u0026ndash;1.37 (Qm\u0026thinsp;=\u0026thinsp;1.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7), broadly ellipsoid, few subglobose and few ellipsoid, inamyloid, hyaline, thin-walled, smooth; apiculus small, 0.5 \u0026micro;m high and 1\u0026ndash;1.5 \u0026micro;m wide. \u003cem\u003ePileipellis\u003c/em\u003e 80\u0026ndash;130 \u0026micro;m thick, 2-layered with radically arranged hyphae; upper layer 30\u0026ndash;35 \u0026micro;m wide, filamentous hyphae gelatinized, few branching, thin-walled, hyaline, 1\u0026ndash;6 \u0026micro;m wide; lower layer 50\u0026ndash;100 \u0026micro;m thick, gelatinized; filamentous hyphae 3\u0026ndash;8 \u0026micro;m wide, hyaline to yellowish. \u003cem\u003eInterior of volva\u003c/em\u003e composed of longitudinally arranged elements: filamentous hyphae 2\u0026ndash;7,5 \u0026micro;m wide, hyaline to yellowish; inflated cells scattered to abundant ellipsoid to elongate 37\u0026ndash;54 \u0026times; 23\u0026ndash;40 \u0026micro;m, hyaline to yellowish, thin-walled. \u003cem\u003eStipe trama\u003c/em\u003e composed of longitudinally arranged, mainly cylindrical, terminal cells, 100\u0026ndash;150 \u0026times; 15\u0026ndash;30 \u0026micro;m; filamentous hyphae scattered, thin-walled, 3\u0026ndash;7 \u0026micro;m wide. \u003cem\u003eClamps\u003c/em\u003e not observed with certainty.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabitat and distribution\u003c/strong\u003e \u003cp\u003eScattered, on soil in mixed highland forests with species of \u003cem\u003ePinaceae\u003c/em\u003e and \u003cem\u003eFagaceae\u003c/em\u003e. Known from China, Loas (this study) Japan, Malaysia and Thailand.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecimens examined\u003c/strong\u003e \u003cp\u003eLaos. Xieng Khouang province, Pek district, Dong village, 19\u0026deg;29'37.8\"N, 103\u0026deg;16'11.2\"E1121 m asl., in mixed forest with \u003cem\u003eQuercus spp.\u003c/em\u003e, \u003cem\u003ePinus kesiya\u003c/em\u003e, and \u003cem\u003eKeteleeria everlyniana\u003c/em\u003e, 14 August 2015, O. Sparre Pedersen and T. L\u0026aelig;ss\u0026oslash;e, OSP20150814-016 (HNL500859); ibid., 19\u0026deg;29'39.8\"N, 103\u0026deg;16'05.8\"E, 1129 m asl., 16 August 2022, O. Sparre Pedersen and W. Sibounheuang, OSP20220816-005 (HNL502458).\u003c/p\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDiscussion\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eAmanita rubromarginata\u003c/em\u003e was described from broad-leaved forests in southwestern islands of Japan (Takahashi \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). It is distinctive by the long striation, i.e. 0.5 R in Cui et al. (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) (in this study 0.4\u0026ndash;0.5 R), the reddish-orange lamellar edge, dark orange to reddish-orange membranous annulus, thick white volva as well as the association with \u003cem\u003eFagaceae\u003c/em\u003e. Notably, we have not seen a greyish-yellow discoloration of aged specimens as shown in Terashima et al. (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Similarly, this feature was similarly not recorded in the description of this species from China (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and from Thailand (Thongbai et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Kumla et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u003cem\u003eAmanita rubromarginata\u003c/em\u003e somewhat resembles \u003cem\u003eA. malayensis\u003c/em\u003e L.P. Tang, Zhu L. Yang \u0026amp; S.S. Lee and \u003cem\u003eA. javanica\u003c/em\u003e (Corner \u0026amp; Bas) T. Oda, C. Tanaka \u0026amp; Tsuda. The former differs by a yellowish to yellowish-orange pileus, concolorous lamellae edge, and the latter differs by less reddish to orange tinges on pileus and annulus. Both of these species are recorded in association with members of \u003cem\u003eDipterocarpaceae\u003c/em\u003e (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, Tulloss and Yang \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). \u003cem\u003eAmanita subhemibapha\u003c/em\u003e Zhu L. Yang, Y.-Y. Cui \u0026amp; Q. Cai (see below) differs by lacking the dark lamellae edge and yellowish zone in lower pileipellis. In our phylogenetic tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) the three lao specimens clustered with strong support the \u003cem\u003eA. rubromarginata\u003c/em\u003e specimens from China and Thailand.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eDescription\u003c/h2\u003e \u003cp\u003e \u003cem\u003eBasidiomata\u003c/em\u003e (Fig.\u0026nbsp;6a, b, e) medium to large sized. \u003cem\u003ePileus\u003c/em\u003e up to 100 mm in diam, initially almost cylindrical, then applanate to slightly depressed, not umbonate, dry or almost so, centre brownish-orange (7C6\u0026ndash;7) when young then reddish-orange (7A8), orange (5A6, 6A8), at margin orange-yellow (4A8); veil remnants absent; margin striate, 0.2\u0026ndash;0.4 R; non-appendiculate; trama white (1A1), unchanging; lower pileipellis with a pale to light yellow (3A3-5) zone, 0.7\u0026ndash;0.9 mm wide. \u003cem\u003eLamellae\u003c/em\u003e free or almost so, yellowish-white (3A2) to light yellow (4A5), 9\u0026ndash;11 mm wide, plentiful (nearly 100 reaching stipe); edge light to pale yellow (3A3\u0026ndash;4); \u003cem\u003elamellulae\u003c/em\u003e truncate, l\u0026ndash;2 shorter of various lengths between full-length lamellae. \u003cem\u003eStipe\u003c/em\u003e 80\u0026ndash;110 \u0026times; 8\u0026ndash;10 mm, cylindrical, hollow, yellowish-white (3A2), partly to fully covered with reddish-orange (7A8) to dark orange (5A8) fibrils; \u003cem\u003evolva\u003c/em\u003e saccate, thin, 35\u0026ndash;60 mm high and 30\u0026ndash;40 mm wide, white (1A1). \u003cem\u003eAnnulus\u003c/em\u003e subapical, membranous, hanging, yellowish-white (3A2) to dark orange (5A8) upper/lower concolorous; upper surface non-striate. \u003cem\u003eTaste\u003c/em\u003e not recorded. \u003cem\u003eSmell\u003c/em\u003e indistinct. \u003cem\u003eSpore deposit\u003c/em\u003e white (1A1).\u003c/p\u003e \u003cp\u003e \u003cem\u003eBasidia\u003c/em\u003e (Fig.\u0026nbsp;6c) 27\u0026ndash;55 \u0026times; 10\u0026ndash;16 \u0026micro;m, clavate, 4-spored; sterigmata 2\u0026ndash;5 \u0026micro;m long. \u003cem\u003eBasidiospores\u003c/em\u003e (Fig.\u0026nbsp;6d) [80/2/2] 7.5\u0026ndash;10 (\u0026ndash;10.5) \u0026times; (6\u0026ndash;) 6.5\u0026ndash;8 (\u0026ndash;8.5) \u0026micro;m (av. 8.5 \u0026times; 7 \u0026micro;m), Q: 1.05\u0026ndash;1.39 (Qm 1.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08) broadly ellipsoid to ellipsoid, few subglobose, inamyloid, hyaline, thin-walled, smooth; apiculus 0.5\u0026ndash;1 \u0026micro;m high and 1\u0026ndash;1.5 \u0026micro;m wide. \u003cem\u003ePileipellis\u003c/em\u003e 120\u0026ndash;175 \u0026micro;m thick; upper layer 50\u0026ndash;100 \u0026micro;m thick, gelatinized, composed of radially to interwoven, thin-walled, hyaline, filamentous hyphae, 2\u0026ndash;3 \u0026micro;m wide; lower layer 90\u0026ndash;120 \u0026micro;m thick, composed of radially to interwoven filamentous hyphae, 4\u0026ndash;15 \u0026micro;m wide, hyaline. \u003cem\u003eInterior of volval remnants\u003c/em\u003e composed of longitudinally to interwoven arranged elements; filamentous hyphae abundant, 1.5\u0026ndash;10 \u0026micro;m wide, hyaline, thin-walled; inflated cells scattered to abundant, broadly ellipsoid, elongate to narrowly cylindrical, hyaline, thin-walled, 38\u0026ndash;125 \u0026times; 30\u0026ndash;60 \u0026micro;m. \u003cem\u003eStipe trama\u003c/em\u003e composed of longitudinally arranged, mainly cylindrical, terminal cells, thin-walled, hyaline, 105\u0026ndash;200 \u0026times; 24\u0026ndash;46 \u0026micro;m; filamentous hyphae scattered, 1.5\u0026ndash;6 \u0026micro;m wide.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabitat and distribution\u003c/strong\u003e \u003cp\u003eScattered on soil in mixed highland forests at altitudes of ca 1150 m, dominated by members of \u003cem\u003ePinaceae\u003c/em\u003e and \u003cem\u003eFagaceae\u003c/em\u003e. Know from Southwestern China, Laos (this study) and northern Thailand.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecimens examined\u003c/strong\u003e \u003cp\u003eLaos. Xieng Khouang province, Pek district, Boa village, 19\u0026deg;29'47.3\"N, 103\u0026deg;17'39.1\"E, 1137 m asl., in mixed forest with \u003cem\u003eCastanopsis spp., Quercus spp.\u003c/em\u003e, few \u003cem\u003ePinus kesiya\u003c/em\u003e and few \u003cem\u003eKeteleeria everlyniana\u003c/em\u003e, 6 August 2017, O. Sparre Pedersen and W. Sibounheuang, OSP20170806-005 (HNL501890); ibid., Dong village, 19\u0026deg;29'32.7\"N, 103\u0026deg;16'06.2\"E, 1141 m asl., in mixed forest with \u003cem\u003eQuercus spp.\u003c/em\u003e, few \u003cem\u003ePinus kesiya\u003c/em\u003e and few \u003cem\u003eKeteleeria everlyniana\u003c/em\u003e, 31 July 2018, O. Sparre Pedersen, OSP20180731\u0026ndash;003 (HNL502059), as \u003cem\u003eA\u003c/em\u003e. cf. \u003cem\u003ehemibapha\u003c/em\u003e in L\u0026aelig;ss\u0026oslash;e et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDiscussion\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eAmanita subhemibapha\u003c/em\u003e was recently described from Yunnan, southwestern China, in association with \u003cem\u003eFagaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e and recognized by the combination of a non-umbonate orange centre of the pileus, striation 0.25\u0026ndash;0.35 R, white to cream lamellae, yellowish lamellae edges and broadly ellipsoid to ellipsoid spores (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The Laotian specimens are slightly darker in the yellow and orange colours; the veil remnants on the stipe are more distinct, and the yellow zone in lower pileipellis is also more pronounced. \u003cem\u003eAmanita subhemibapha\u003c/em\u003e somewhat resembles \u003cem\u003eA. hemibapha\u003c/em\u003e (Berk. \u0026amp; Broome) Sacc., \u003cem\u003eA. malayensis, A. kitamagotake\u003c/em\u003e N. Endo \u0026amp; A. Yamada, \u003cem\u003eA. rubromarginata\u003c/em\u003e and \u003cem\u003eA. javanica. Amanita hemibapha\u003c/em\u003e differs by having a darker red pileus centre, an orange margin and narrower spores (5.5\u0026ndash;6.5 \u0026micro;m) (Vrinda et al \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) to more yellowish pileus margin and spore width of 5.5\u0026ndash;7 \u0026micro;m (Kumla et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u003cem\u003eAmanita rubromarginata\u003c/em\u003e and \u003cem\u003eA. javanica\u003c/em\u003e differ by having longer pileal striation (0.4\u0026ndash;0.5 R) and reddish lamellae edges; \u003cem\u003eA. kitamagotake\u003c/em\u003e differs by having uniformly yellow colours on pileus and stipe surface (Yang \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Endo et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e)d \u003cem\u003emalayensis\u003c/em\u003e differs by having a whitish-cream stipe and an association with dipterocarps (Tang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Our phylogenetic tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) indicates that \u003cem\u003eA. subhemibapha\u003c/em\u003e is closely related to \u003cem\u003eA. malayensis A. similis\u003c/em\u003e Boedijn and \u003cem\u003eA. fuscoflava\u003c/em\u003e, Zhu L. Yang, Yang-Yang Cui \u0026amp; Qing Cai. The former differences to \u003cem\u003eA. subhemibapha\u003c/em\u003e are discussed above and under the discussion heading of \u003cem\u003eA. rubromarginata; A. similis\u003c/em\u003e differs by a longer pileal striation (0.33\u0026ndash;0.6 R) and a greyish brown to olivaceous brown pileal center (Boedijn (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1951\u003c/span\u003e; Corner and Bas \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1962\u003c/span\u003e); and \u003cem\u003eA. fuscoflava\u003c/em\u003e also differs by a longer pilar striation (0.5\u0026ndash;0.6 R) and a dark brown pilar center (Cui et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eKey to\u003c/b\u003e \u003cb\u003eAmanita\u003c/b\u003e \u003cb\u003esect.\u003c/b\u003e \u003cb\u003eCaesareae\u003c/b\u003e \u003cb\u003eknown from and likely to occur in Laos\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003col\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003eStipe white to cream with fibrils of same colours 2\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003c/ol\u003e \u003c/p\u003e \u003cp\u003eStipe yellowish-white to yellowish, typically with darker fibrils 8\u003c/p\u003e \u003cp\u003e2. Basidiomata entirely white or cream to alutaceous-cream at centre of pileus 3\u003c/p\u003e \u003cp\u003eBasidiomata with yellow-greyish, cream, brown to brownish centre of pileus 5\u003c/p\u003e \u003cp\u003e3. Striation on pileus 0.3\u0026ndash;0.4 R; annulus persistent. Pileus yellowish with age; pileus centre cream; stipe white; basidia with 4 sterigmata; spores globose to sub-globose, 9\u0026ndash;11 \u0026times; 9\u0026ndash;11 \u0026micro;m, Q\u0026thinsp;=\u0026thinsp;1.0\u0026ndash;1.13, Qm\u0026thinsp;=\u0026thinsp;1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 \u003cem\u003eA. alboumbelliformis\u003c/em\u003e\u003c/p\u003e \u003cp\u003eStriation on pileus 0.1\u0026ndash;0.3 R; annulus fragile 4\u003c/p\u003e \u003cp\u003e4. Pileus centre cream to alutaceous cream, striation 0.25 R. Annulus white; stipe squamules white in snakeskin pattern, basidia mainly with 2 sterigmata; spores globose to subglobose, 8.5\u0026ndash;10.5 \u0026times; 8.5\u0026ndash;9.5 \u0026micro;m, Q\u0026thinsp;=\u0026thinsp;1.04\u0026ndash;1.14, Qm\u0026thinsp;=\u0026thinsp;1.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 \u003cem\u003eA.\u003c/em\u003e \u003cem\u003echepangianoides\u003c/em\u003e\u003c/p\u003e \u003cp\u003ePileus centre white, sometimes with yellowish, yellowish or brownish tint, striation 0.1\u0026ndash;0.15R. Annulus with age yellowish; stipe squamules white, bruising yellowish to greyish; basidia with 4 sterigmata; spores sub- globose to broadly ellipsoid, 9.5\u0026ndash;12.5 \u0026times; 8.5\u0026ndash;11.5 \u0026micro;m, Q\u0026thinsp;=\u0026thinsp;1.07\u0026ndash;1.28, Qm\u0026thinsp;=\u0026thinsp;1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 \u003cem\u003eA\u003c/em\u003e. \u003cem\u003echepangiana\u003c/em\u003e\u003c/p\u003e \u003cp\u003e5. Striation on pileus 0.3\u0026ndash;0.6 R; spores globose to subglobose 6\u003c/p\u003e \u003cp\u003eStriation on pileus less than 0.3 R; spores subglobose to broadly ellipsoid 7\u003c/p\u003e \u003cp\u003e6. Pileus to 140 mm wide, with broad umbo, striation 0.3\u0026ndash;0.4 R, centre first greyish-orange soon golden brown. Stipe white, smooth to slightly fibrous; volval surface not cracking into brownish patches; annulus membranous, basidiospores 9\u0026ndash;12 \u0026times; 8.5\u0026ndash;11.5 \u0026micro;m, Qm: 1.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 \u003cem\u003eA. fulvocaesarea\u003c/em\u003e\u003c/p\u003e \u003cp\u003ePileus 60\u0026ndash;100 mm wide, slightly depressed, non-umbonate, striation 0.5\u0026ndash;0.6 R, centre reddish-brown.\u003c/p\u003e \u003cp\u003eStipe pale cream, \u0026plusmn; darker squamules; anulus fragile; basidiospores 8.5\u0026ndash;11 \u0026times; 8.5\u0026ndash;10.5 \u0026micro;m, Qm: 1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 \u003cem\u003eA. aporema\u003c/em\u003e\u003c/p\u003e \u003cp\u003e7. Pileus to 150 mm in diameter, centre yellowish cream, olive-grey to yellowish brown; stipe squamules white to dirty white; volval surface not cracking into brownish patches; basidiospores 9.5\u0026ndash;14.5 \u0026times; 8\u0026ndash;12.5 \u0026micro;m, Qm\u0026thinsp;=\u0026thinsp;1.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 \u003cem\u003eA. pseudoprinceps\u003c/em\u003e\u003c/p\u003e \u003cp\u003ePileus to 200 mm in diameter, centre brown to yellowish-brown; stipe squamules white, volva often cracking and peeling into thin patches; basidiospores 8\u0026ndash;11 \u0026times; 7.5\u0026ndash;10 \u0026micro;m, Qm\u0026thinsp;=\u0026thinsp;1.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 \u003cem\u003eA. princeps\u003c/em\u003e\u003c/p\u003e \u003cp\u003e8. Lamellar edge concolorous with lamellae 9\u003c/p\u003e \u003cp\u003eLamellar edge yellowish or reddish orange 11\u003c/p\u003e \u003cp\u003e9. Pileus uniformly yellowish. Pileus 40\u0026ndash;100 mm in diameter, striation 0.3\u0026ndash;0.5 R; lamellae yellowish; stipe yellowish to yellow, squamules yellow to orange buff; annulus yellow; basidiospores broadly ellipsoid to ellipsoid, 8.5\u0026ndash;13.5 \u0026times; 6\u0026ndash;8.5 \u0026micro;m, Qm\u0026thinsp;=\u0026thinsp;1.41\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6 \u003cem\u003eA. kitamagotake\u003c/em\u003e\u003c/p\u003e \u003cp\u003ePileus centre and margin of different colours 10\u003c/p\u003e \u003cp\u003e10. Pileus centre red to orange red to yellow, margin yellow. Pileus 60\u0026ndash;150 mm in diameter, striation 0.3\u0026ndash;0.5R;\u003c/p\u003e \u003cp\u003elamellae cream to yellowish; stipe yellowish, stipe squamules yellow to orange-yellow in irregular snakeskin pattern; annulus yellow; basidiospores ellipsoid, 7.5\u0026ndash;11 \u0026times; 5.5\u0026ndash;7.5 \u0026micro;m, Qm\u0026thinsp;=\u0026thinsp;1.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 \u003cem\u003eA. hemibapha\u003c/em\u003e\u003c/p\u003e \u003cp\u003ePileus centre yellowish to yellowish orange. Pileus 60\u0026ndash;100 mm in diameter, margin pale yellow to maize yellow, striation 0.3\u0026ndash;0.4R; stipe yellowish to cream, stipe squamules below annulus yellow; lamellae whitish, edge colour unknown probably concolorous; annulus yellowish white to whitish; basidiospores broadly ellipsoid to ellipsoid, 9\u0026ndash;12 \u0026times; 6\u0026ndash;8 \u0026micro;m, Qm 1.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 \u003cem\u003eA. malayensis\u003c/em\u003e\u003c/p\u003e \u003cp\u003e11. Lamellar edge reddish orange. Pileus center orange red to brownish-red to red, margin orange to deep yellow, 60\u0026ndash;90(130) mm in diameter; striation 0.4\u0026ndash;0.5 R; stipe yellowish, stipe squamules orange to orange-red, often in a snakeskin pattern; lamellae yellow, edge light orange to reddish-orange; annulus, dark orange to reddish-orange; basidiospores 6.5\u0026ndash;9 \u0026times; 5.5\u0026ndash;7 \u0026micro;m, Qm 1.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07. \u003cem\u003eA. rubromarginata\u003c/em\u003e\u003c/p\u003e \u003cp\u003eLamellar edge in yellow colours 12\u003c/p\u003e \u003cp\u003e12. Pileus center brownish-orange to reddish-orange; margin orange yellow, 60\u0026ndash;110 mm in diameter; striation 0.2\u0026ndash;0.35 R. Lower pileipellis with yellowish zone (0.7\u0026ndash;0.9 mm); stipe yellowish white, stipe squamules reddish-orange to dark orange; lamellae yellowish-white to light yellow, annulus yellowish-white to orange; basidiospores 7.5\u0026ndash;10 \u0026times; 6.5\u0026ndash;8 \u0026micro;m, Qm: 1.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 \u003cem\u003eA. subhemibapba\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003ePileus center orange-yellow to ochre yellow, margin paler or buff, 60\u0026ndash;90 mm in diameter; striation 0.4\u0026ndash;0.5 R. Lamellae whitish, stipe yellowish, stipe squamules orange-buff; annulus orange buff, membranous;\u003c/p\u003e \u003cp\u003ebasidiospores 7.5\u0026ndash;9 \u0026times; 5.8\u0026ndash;7 \u0026micro;m, Qm: 1.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 \u003cem\u003eA. javanica\u003c/em\u003e\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUrmas K\u0026otilde;ljalg, Irja Saar and staff at Institute of Ecology and Earth Sciences, Department of Plant Pathology, University of Tartu, Estonia, are gratefully thanked for carrying out DNA analyses and uploading to the PlutoF platform. The encouragements from and many discussions with Andy Taylor of the James Hutton Institute are sincerely appreciated. Finally, Yang-Yang Cui and Zhu Liang Yang are heartfully thanked for their constructive suggestions and for allowing us to use unpublished data.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eField work in Laos was partly GEF-financed under the UNDP-FAO Agrobiodiversity Project (UNTS/LAO/015/GEF), partly by the Darwin Initiative Project \u0026ldquo;Building mycological capacity for sustainable resources management in Lao PDR\u0026rdquo; of The James Hutton Institute and partly through private means.\u003c/p\u003e\n\u003cp\u003eDrafts of the manuscript were written by OSP with regular comments and suggestions from TL. Both authors have read and approved the final manuscript. \u003cem\u003eDatasets generated and analyzed during the current study are available from the corresponding author on reasonable request.\u0026nbsp;\u003c/em\u003eThe authors declare no competing interests related to this study.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbarenkov K, Nilsson RH, Larsson K-H, Taylor AFS, May TW, Fr\u0026oslash;slev TG, Pawlowska J, Lindahl B, P\u0026otilde;ldmaa K, Truong C, Vu D, Hosoya T, Niskanen T, Piirmann T, Ivanov F, Zirk A, Peterson M, Cheeke TE, Ishigami Y, Jansson AT, Jeppesen TS, Kristiansson E, Mikryukov V, Miller JT, Oono R, Ossandon FJ, Paup\u0026eacute;rio J, Saar I, Schigel D, Suija A, Tedersoo L, K\u0026otilde;ljalg U (2023) The UNITE database for molecular identification and taxonomic communication of fungi and other eukaryotes: sequences, taxa and classifications reconsidered. 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Environmentalist 8:187\u0026ndash;208 \u003c/li\u003e\n\u003cli\u003eNAFRI, NUoL, SNV (2007) Non-timber forest products in the Lao PDR: a manual of 100 commercial and traditional products, National Agriculture and Forestry Institute, Vientiane\u003c/li\u003e\n\u003cli\u003eNewman M, Ketphanh S, Svengsuksa, B, Thomas P, Sengdala K, Lamxay V, Amstrong V (2007) A checklist of vascular plants of Lao PDR, Royal Botanic Garden, Edinburgh. https://portals.iucn.org/library/efiles/documents/2007-014.pdf\u003c/li\u003e\n\u003cli\u003ePatouillard NT (1928) Nouvelle contribution \u0026agrave; la flore mycologique de l\u0026apos;Annam et du Laos. Annales de Cryptogamie Exotique 1: 2\u0026ndash;24. https://bibdigital.rjb.csic.es/medias/9b/7c/a3/72/9b7ca372-b7fb-405c-bf57-4411ce78fe06/files/P1661_01.pdf\u003c/li\u003e\n\u003cli\u003ePedersen OS, Thammavong K (2014) Biodiversity, Wild Mushrooms, edible and medicinal species, local knowledge and use. Pilot survey. 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The Victorian Naturalist. 95:47\u0026ndash;49. https://www.biodiversitylibrary.org/item/123295#page/54/mode/1up\u003c/li\u003e\n\u003cli\u003eRonquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. 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Current Research in Environmental \u0026amp; Applied Mycology (Journal of Fungal Biology) 12(1): 65\u0026ndash;74. doi 10.5943/cream/12/1/6\u003c/li\u003e\n\u003cli\u003eSysouphanthong P, Thongklang N, Liu Y, Vellinga E. (2022) Three new species of \u003cem\u003eCystolepiota\u003c/em\u003e from Laos and Thailand. Diversity14(6):449. https://doi.org/10.3390/d14060449\u003c/li\u003e\n\u003cli\u003eTakahashi H (2004) Two new species of Agaricales from southwestern islands of Japan, Mycoscience 45(6):372\u003c/li\u003e\n\u003cli\u003eTamura K, Stecher G, Kumar S. MEGA11 (2021) Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution. 38(7): 3022\u0026ndash;27. https://doi.org/10.1093/molbev/msab120\u003c/li\u003e\n\u003cli\u003eTang L-P, Lee S-S, Zeng N-K, Cai Q, Zhang P, Zhu L, Yang ZL (2017) Notes on \u003cem\u003eAmanita \u003c/em\u003esection \u003cem\u003eCaesareae\u003c/em\u003e from Malaysia. 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J Ethnobiol Ethnomed 18:55. https://doi.org/10.1186/s13002-022-00551-7\u003c/li\u003e\n\u003cli\u003eYang ZL (2015) Atlas of the Chinese species of \u003cem\u003eAmanitaceae\u003c/em\u003e [In Chinese]. ISBN 9787030436276, Science Press, Beijing \u003c/li\u003e\n\u003cli\u003eYang ZL, Cai Q, Cui, YY (2018) Phylogeny, diversity and morphological evolution of \u003cem\u003eAmanitaceae\u003c/em\u003e. Biosyst Ecol Ser 34:359\u0026ndash;380\u003c/li\u003e\n\u003c/ol\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":"Amanitaceae, taxonomy, South–East Asia, Fagaceae, Pinaceae, 2 new taxa","lastPublishedDoi":"10.21203/rs.3.rs-4401254/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4401254/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSpecimens of \u003cem\u003eAmanita\u003c/em\u003e sect. \u003cem\u003eCaesareae\u003c/em\u003e, collected in Xieng Khouang and Vientiane provinces in Laos from highland forests dominated by members of \u003cem\u003eFagaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e, are studied based on morphological characters and analyses of ITS sequences. Two species, namely \u003cem\u003eA. fulvocaesarea\u003c/em\u003e and \u003cem\u003eA. chepangianoides\u003c/em\u003e are reported as new, both found in association with \u003cem\u003eFagaceae\u003c/em\u003e, whereas \u003cem\u003eA. pseudoprinceps and A. subhemibapha\u003c/em\u003e, recently described from China and \u003cem\u003eA. rubromarginata\u003c/em\u003e from Japan, are confirmed from Laos and found in forests with \u003cem\u003eFagaceae\u003c/em\u003e and \u003cem\u003ePinaceae\u003c/em\u003e. The phylogenetic analyses to recognize the \u003cem\u003eAmanita\u003c/em\u003e species showed that the golden-brown to greyish orange \u003cem\u003eA. fulvocaesarea\u003c/em\u003e and the dominantly white \u003cem\u003eA. chepangianoides\u003c/em\u003e cluster with \u003cem\u003eA. aporema, A. chepangiana, A. egregia\u003c/em\u003e, \u003cem\u003eA. princeps\u003c/em\u003e and \u003cem\u003eA. pseudoprinceps.\u003c/em\u003e A dichotomous key to identified and likely occurring \u003cem\u003eAmanita\u003c/em\u003e species in Laos within sect. \u003cem\u003eCaesareae\u003c/em\u003e is also provided.\u003c/p\u003e","manuscriptTitle":"Amanita species within section Caesareae from the highlands of Laos","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-05-29 14:57:30","doi":"10.21203/rs.3.rs-4401254/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2024-05-18T04:05:09+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-18T04:03:03+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Mycological Progress","date":"2024-05-15T14:53:40+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-05-13T07:39:41+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mycological Progress","date":"2024-05-10T10:20:11+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":"a61c8f32-905d-469b-91ee-9a72cf37f643","owner":[],"postedDate":"May 29th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-10-07T16:08:32+00:00","versionOfRecord":{"articleIdentity":"rs-4401254","link":"https://doi.org/10.1007/s11557-024-01999-5","journal":{"identity":"mycological-progress","isVorOnly":false,"title":"Mycological Progress"},"publishedOn":"2024-10-05 15:58:00","publishedOnDateReadable":"October 5th, 2024"},"versionCreatedAt":"2024-05-29 14:57:30","video":"","vorDoi":"10.1007/s11557-024-01999-5","vorDoiUrl":"https://doi.org/10.1007/s11557-024-01999-5","workflowStages":[]},"version":"v1","identity":"rs-4401254","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4401254","identity":"rs-4401254","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}