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Article

Taxonomic Novelties and New Records of Amanita Subgenus Amanitina from Thailand

by
Yuan S. Liu
1,2,3,
Jian-Kui Liu
4,
Jaturong Kumla
1,3,
Nakarin Suwannarach
1,3 and
Saisamorn Lumyong
1,3,5,*
1
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
2
Doctor of Philosophy Program in Applied Microbiology (International Program), Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
3
Research Center of Microbial Diversity and Sustainable Utilization, Chiang Mai University, Chiang Mai 50200, Thailand
4
School of Life Science and Technology, Center for Informational Biology, University of Electronic Science and Technology of China, Chengdu 611731, China
5
Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand
*
Author to whom correspondence should be addressed.
J. Fungi 2023, 9(6), 601; https://doi.org/10.3390/jof9060601
Submission received: 9 April 2023 / Revised: 16 May 2023 / Accepted: 17 May 2023 / Published: 24 May 2023
(This article belongs to the Special Issue Recent Advances in Taxonomy, Phylogeny and Evolution of Fungi)
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Abstract

:
The Amanita subgenus Amanitina contains six sections, and the species diversity of this subgenus has still not been explored in Thailand. Twenty samples collected in 2019 and 2020, which had the morphological characteristics of the Amanita subgen. Amanitina, were observed in this study. Both the microscopical characteristics and multi-gene phylogenetic analyses of the ITS, nrLSU, RPB2, TEF1-α, and TUB gene regions revealed that the 20 samples represented nine species and dispersed into four sections. Remarkably, three taxa were different from any other currently known species. Here, we describe them as new to science, namely A. albifragilis, A. claristriata, and A. fulvisquamea. Moreover, we also recognized six interesting taxa, including four records that were new to Thailand, viz. A. cacaina, A. citrinoannulata, A. griseofarinosa, and A. neoovoidea, as well as two previously recorded species, A. caojizong and A. oberwinkleriana. Moreover, we provide the first RPB2 and TEF1-α gene sequences for A. cacaina. Detailed descriptions, illustrations as line drawings, and comparisons with related taxa are provided.

1. Introduction

Amanita Pers. is an important basidiomycetous genus comprising about 700 species [1,2,3,4]. It contains both well-known edible and deadly poisonous species. In addition, Amanita species are regarded as key organisms involved in nutrient and carbon cycling in forest ecosystems on account of their ability to form ectomycorrhizal relationships with more than 10 families of vascular plants, e.g., Dipterocarpaceae, Fagaceae, Myrtaceae, and Pinaceae [2,5,6,7].
Since the genus Amanita was formally established in 1797 [8], many mycologists have continued to contribute to and improve the taxonomic knowledge of this genus [9,10,11,12,13,14,15,16,17,18,19,20,21,22]. Corner and Bas [15] and Bas [16] proposed splitting the genus Amanita into two subgenera and six sections, which had important significance for the taxonomy of Amanita. Yang [17] revised the classification of this genus and split it into two subgenera and seven sections. Although the above classifications have been widely adopted, the delimitation within this genus is still controversial [23,24]. Until 2018, according to multi-gene phylogenetic analysis, morphological examinations, and ecological studies, Cui et al. [2] proposed the division of the genus Amanita into three subgenera and eleven sections as follows. The subgenus Amanita contains the section Amanita, section Amarrendiae (Bougher and T. Lebel) Zhu L. Yang, Y.Y. Cui, Q. Cai, and L.P. Tang, section Caesareae Singer ex Singer, and section Vaginatae (Fr.) Quél. The subgenus Amanitina (E. J. Gilbert) E.J. Gilbert contains the section Amidella (J. E. Gilbert) Konrad and Maubl., section Arenariae Zhu L. Yang, Y.Y. Cui and Q. Cai, section Phalloideae (Fr.) Quél., section Roanokenses Singer ex Singer, section Strobiliformes Singer ex Q. Cai, Zhu L. Yang and Y.Y. Cui, and section Validae (Fr.) Quél. The subgenus Lepidella Beauseigneur contains the section Lepidella Corner and Bas. Under this treatment, all saprotrophic Amanita species are assigned to the section Lepidella.
Surveys of species diversity, which can provide abundant materials, are fundamental and important work within mycological research. As more Amanita species are found and documented, researchers can have a better understanding and knowledge of this genus. However, the level of knowledge on the species diversity of Thai Amanita remains limited [25,26,27,28]. Through studying materials collected by their group over an extensive period and borrowed from other herbaria, Sanmee et al. [25] comprehensively reported 25 taxa of Thai Amanita with formal and detailed descriptions, including 18 records that were new to Thailand and seven known species. Between 2016 and 2018, Thongbai et al. [26,29,30] originally reported 12 species based on both morphology and phylogeny, which made an important contribution to the taxonomy of Thai Amanita. In addition, a number of researchers have worked on the species diversity of Amanita in Thailand [27,28,31,32,33,34,35], including those from our research group [33,34,35]. Up to now, 59 taxa have been reported from Thailand. Among these species, 21 were first described from Thailand and 24 belonged to the subgenus Amanitina according to the latest classification [25,26,28,29,30,31,32,33,34,35].
In the present study, we examined 20 specimens, which were collected from deciduous or coniferous forests predominantly composed of Dipterocarpus, Shorea, or Pinus species in northern and northeastern Thailand. On the basis of the macro- and microscopical characteristics, as well as multi-gene phylogenetic analyses, we identified them as nine Amanita species classified in the sections Amidella, Phalloideae, Roanokenses, and Validae. Among these taxa, three are reported as new to science, and four are, for the first time, reported from Thailand.

2. Materials and Methods

2.1. Morphological Study

Basidiomata was collected from deciduous or coniferous forests in Chiang Mai, Chiang Rai, Phetchabun, and Sakon Nakho provinces in Thailand during the rainy season of 2019 and 2020. The following information was recorded: color photographs, forest type, substrate type, and geographic coordinates. Small pieces of tissue from the cap and/or stipe were taken and dried with silica gel to prepare for the molecular analyses [36], and the remaining specimens were dried at 35–45 °C for at least 12 h to prepare for the morphological examinations. All specimens observed in this study were deposited at the Herbarium of Biology Department (CMUB) and the Herbarium of Sustainable Development of Biological Resources (SDBR), Faculty of Science, Chiang Mai University, Thailand.
The macroscopic characteristics were described on the basis of field notes and images. The color codes and names were described according to Kornerup and Wanscher [37]. The microscopic features were observed in distilled water, 5% aqueous KOH (w/v), 1% Congo red (w/v), and Melzer’s reagent under a Leica DM500 microscope [18,19]. Sections of the pileipellis were cut along radial planes taken from halfway between the center and the margin of the pileus. Sections of the stipitipellis were cut longitudinally from small pieces taken from the middle part of the stipe. For the description of the basidiospores, the term (n/m/p) represents n basidiospores measured from m basidiomata of p collections. The dimensions for the basidiospores are given as (a–) b–c (–d), in which ‘b–c’ represents a minimum of 90% of the measured and extreme values, and ‘a’ and ‘d’ are given in parentheses whenever necessary. Q denotes the ratio of the length divided by the width of the basidiospore from the side view, Qm denotes the average Q of n measured basidiospores, and SD is their standard deviation. The results are presented as Q = Qm ± SD. Marginal striations on the pileus are expressed as a proportion of the ratio of the length of the striation to the radius of the pileus (nR). The terms denoting the basidiomata size and the spore shape are defined according to Bas [16] and Yang [19].

2.2. DNA Extraction, PCR Amplification, and Sequencing

Detailed processes of DNA extraction, PCR amplification, and sequencing protocols were carried out in line with previous studies [33,38]. Five DNA gene fragments were amplified and sequenced, including the internal transcribed spacer region (ITS), the large subunit of the nuclear ribosomal DNA (nrLSU), the partial sequences of the RNA polymerase II second largest subunit (RPB2), the translation elongation factor 1-alpha (TEF1-α), and the beta-tubulin gene (TUB).

2.3. Phylogenetic Analyses

The newly generated sequences were used for BLAST searching in NCBI GenBank (https://www.ncbi.nlm.nih.gov, accessed on 7 March 2023), and then closely related sequences and Thai sequences were selected for the initial analysis. Detailed information, including the newly generated sequences and the sequences obtained from GenBank, is provided in Table 1.
Sequences of each gene fragment were separately aligned with MAFFT v.7 [39] using the G-INSi iterative refinement algorithm and then manually optimized with AliView v.1.28 [40]. Gblocks v.0.91b [41] was used to exclude the ambiguously aligned regions for ITS with two options: “Allow smaller final blocks” and “Allow gap positions within the final blocks”. Sequence Matrix v.100.0 was applied to concatenate the five gene fragments for further phylogenetic analysis. MrModeltest v.2.3 [42] was adopted to determine the best fitting model of nucleotide substitution for each single-gene dataset by applying the default parameters.
Phylogenetic trees were inferred using both maximum likelihood (ML) and Bayesian inference (BI), as detailed in [43]. The ML analysis was performed at the CIPRES web portal [44] using RAxML v.8.2.12 as part of the “RAxML-HPC BlackBox” tool [45] with the default settings, and the option “Estimate proportion of invariable sites (GTRGAMMA+I)” was set to “yes” for both the single-gene and the concatenated gene analyses. The phylogenetic analyses were initially performed on each single-gene alignment, and since there was no evident conflict (with ML bootstrap support of ≥75%), the concatenated dataset was built, and the multi-gene ML analysis was performed. The Bayesian analysis was carried out with MrBayes v.3.1.2 [46]. The posterior probabilities [47] were determined via Markov chain Monte Carlo sampling (MCMC) [48]. Six simultaneous Markov chains were run from random trees for one million generations, and the trees were sampled every 100th generation (the critical value for diagnosing topological convergence was 0.01). The first 25% of the trees were discarded, and the remaining trees were used for calculating the posterior probabilities in the majority-rule consensus tree. The phylogenetic trees were visualized with FigTree v.1.4.4 [49].

3. Results

3.1. Phylogenetic Analyses

The best fitting model for each gene fragment was as follows: general time reversible + proportion of invariable sites + gamma distribution (GTR + I + G) for nrLSU, RPB2, TEF1-α, and TUB; Hasegawa–Kishino–Yano (HKY) + I + G for ITS. The concatenated dataset was partitioned into five parts according to the sequence region. Because the model HKY + I + G could not be implemented in ML, the GTR + I + G model was used, as it included all the parameters of the selected model.
The multi-gene dataset comprised 466 sequences, of which 71 were newly generated and 395 were retrieved from GenBank. Amanita flavofloccosa (HKAS101443), A. flavofloccosa (HKAS90174), and A. vittadinii (HKAS101430) from Amanita section Lepidella were chosen as the outgroup taxa. The final multi-gene alignment comprised 3650 positions (nrLSU: 1–944; ITS: 945–2002; RPB2: 2003–2676; TEF1-α: 2677–3254; and TUB: 3255–3650), including gaps.
The resulting topologies of the ML and BI analyses were congruent; therefore, an ML tree is shown in Figure 1. In our phylogenetic analyses, all six sections of the subgenus Amanitina showed similar mutual relationships as those in previous studies [2,4], as well as the species in each section. The three novel species formed a clearly monophyletic lineage that was distinct from other extant species with credibly supported values.

3.2. Taxonomy

Amanita sect. Amidella (E. J. Gilbert) Konrad and Maubl., Agaricales: 61 (1948).
Basionym: Amidella E. J. Gilbert, in Bres., Iconogr. Mycol. 27 Suppl. 1(1): 71 (1940).
Type: Amanita volvata (Peck) Lloyd, Mycol. Writ. 1(7): 9 (1898).
Notes: Species from section Amidella have a series of remarkable characteristics, such as the color of the basidiomata changing to a brownish or reddish tone when injured, a striate and appendiculate pileal margin, the lamellae changing to a brown tone upon drying, truncate lamellulae, the amyloid basidiospores, and the absence of clamps [2,16,50]. The above combination of characteristics is unique to the section Amidella and is not found in any other section of Amanita.
Presently, only two species of the section Amidella have been reported from Thailand, namely Amanita avellaneosquamosa (S. Imai) S. Imai and A. clarisquamosa (S. Imai) S. Imai [25]. In this study, six specimens collected from Chiang Mai and Phetchabun provinces were recognized and described as two novel species belonging to the section Amidella.
Amanita claristriata Yuan S. Liu and S. Lumyong, sp. nov.; Figure 2a and Figure 3.
MycoBank number: 847954
Etymology: “claristriata”, from clarus (obvious) and striatus (grooved), indicates that this species has obvious striations on the margin of its pileus.
Holotype: THAILAND, Chiang Mai Province, Mueang District, 18°48′24.3″ N 98°54′38.1″ E, alt. 1102 m, 3 September 2020, Yuan S. Liu, STO-2020-404 (CMUB39992). GenBank accession numbers: OQ780686 (ITS), OQ780668 (nrLSU), OQ740048 (RPB2), and OQ740066 (TEF1-α).
Basidiomata medium-sized. Pileus 5.6–7.2 cm in diam., plano-convex to applanate, sometimes depressed at the center, white (1A1) to orange white (6A2); volval remnants on the pileus floccose to scaly, sometimes disappear because of rain, greyish orange (6B5–6) to brownish orange (7C5–6), densely arranged over the disk; margin inconspicuously striate at first and becoming obviously so with age, sometimes up to 0.3 R, appendiculate; context 5.0–7.0 mm wide, white (1A1), changing to orange white or pale orange (6A2–3) after injury. Lamellae free, crowded, white (1A1), becoming brown to dark brown (6F5–8) upon drying; lamellulae mostly truncate. Stipe 11.3–16.0 cm long × 0.7–1.3 cm diam. (the length includes the basal bulb), subcylindric or slightly tapering upwards, with the apex slightly expanded, white (1A1), covered with fibrous to floccose, white (1A1), greyish orange (6B3–4) to brown (6C4–6) squamules; context white (1A1), changing to orange white or pale orange (6A2–3) after injury, fistulose; basal bulb absent; volva saccate, 3.0–4.2 cm high × 1.8–2.4 cm wide, membranous, white (1A1) to greyish orange (6B5–6). Annulus absent. Odor not recorded.
Lamellar trama bilateral. Mediostratum 30–40 μm wide, consisting of abundant clavate inflated cells (45–115 × 10–20 μm); filamentous hyphae abundant, 2–8 μm wide; vascular hyphae scarce. Lateral stratum 30–40 μm thick, consisting of abundant to dominated clavate inflated cells (30–72 × 9–16 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 2–6 μm wide. Subhymenium 20–30 μm thick, with two–three layers of subglobose or irregular cells, 9–23 × 6–16 μm. Basidia (Figure 3b) 33–47 × 8–11 μm, clavate, four-spored; sterigmata 3–6 μm long; basal septa lacking clamps. Basidiospores [82/3/3] (8.0–) 9.0–11.5 × 5.0–6.0 μm, avl × avw = 10.3 × 5.3 μm, Q = (1.60–) 1.64–2.20 (–2.30), Qm = 1.95 ± 0.18, elongate to cylindrical, thin-walled, smooth, colorless to pastel yellow, amyloid (Figure 3a). Lamellar edge sterile, consisting of subglobose to ellipsoid or clavate inflated cells (13–45 × 8–23 μm), single or in chains of 2–3, thin-walled, colorless to pastel yellow; filamentous hyphae abundant, 3–6 μm wide, irregularly arranged. Pileipellis 70–140 μm thick, two-layered; upper layer (30–80 μm thick) gelatinized, consisting of radially, thin-walled, colorless to pale yellow, filamentous hyphae 1–4 μm wide; lower layer (35–65 μm thick) consisting of radially and compactly arranged, filamentous hyphae 2–8 μm wide, colorless to pale yellow; vascular hyphae scarce. Volval remnants on pileus (Figure 3c) consisting of subradially to radially arranged elements: filamentous hyphae abundant, 2–10 μm wide, colorless pale yellow, thin-walled; inflated cells abundant, ellipsoid to clavate, 60–270 × 20–37 μm, colorless to pale yellow, thin-walled, often terminal; vascular hyphae scarce. Interior of volval remnants on stipe base consisting of sub-longitudinally to longitudinally arranged elements: filamentous hyphae abundant, 2–10 μm wide, colorless, thin-walled; inflated cells fairly abundant, subglobose to ellipsoid or ovoid, 40–98 × 15–55 μm, colorless, thin-walled, often terminal; vascular hyphae scarce. The outer surface of volval remnants on stipe base consisting of very abundant filamentous hyphae (2–6 μm wide), mixed with scattered to fairly abundant, subglobose to ellipsoid, or ovoid inflated cells. The inner surface of volval remnants on stipe base gelatinized, similar to structure of interior part but comprising much more filamentous hyphae (1–5 μm wide). Stipe trama consisting of longitudinally arranged, long clavate, terminal cells, 125–365 × 20–45 μm; filamentous hyphae scattered to abundant, 2–5 μm wide; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: Currently known in northern Thailand.
Additional collections examined: THAILAND, Chiang Mai Province, Mueang District, alt. 1102 m, 3 September 2020, Yuan S. Liu, STO-2020-407 (SDBR-STO20-407); Yuan S. Liu, STO-2020-408 (SDBR-STO20-408).
Notes: Amanita claristriata is characterized by its medium-sized basidiomata, orange white pileus covered by floccose-felted to patchy, brownish orange volval remnants, an obvious striate margin on pileus (after maturity), the color of the basidiomata changes when injured (from white to pale orange), longer stipe (11.3–16.0 cm) covered by fibrous to floccose, brownish squamules, saccate volva remnants on the stipe base, as well as elongate to cylindrical basidiospores (9.0–11.5 × 5.0–6.0 μm, Qm = 1.95 ± 0.18).
Morphologically, Amanita lanigera Y.Y. Cui, Q. Cai and Zhu L. Yang and A. parvicurta Y.Y. Cui, Q. Cai and Zhu L. Yang resemble A. claristriata. Amanita lanigera, described from China, differs from A. claristriata by having the non-striate pileal margin (or slightly striate), basidiomata color unchanged when injured, and larger, ellipsoid, colorless basidiospores (10.0–12.0 × 7.0–8.5 μm, Qm = 1.49 ± 0.13) [2].
Moreover, it is remarkable that Amanita rufobrunnescens W. Q. Deng and T. H. Li [51] and A. volvata [16,52,53] share a particular and consistent feature with A. claristriata, viz. the basidiomata changes to light red or pale orange after injury. However, both A. rufobrunnescens reported from China and A. volvata reported from America have larger basidiospores (10.0–12.0 × 5.5–6.5 μm, Qm = 1.78 ± 0.17 for A. rufobrunnescens; 10.0–12.5 × 6.0–7.5 μm, Qm = 1.67 ± 0.11 for A. volvata) [2,16,51,53].
Phylogenetically, Amanita claristriata is related to A. peckiana Kauffman and A. pinophila Y.Y. Cui, Q. Cai and Zhu L. Yang. However, the latter two species differ from the former by not changing basidiomata color when injured, as well as larger basidiospores (9.8–13.6 × 5.6–7.0 μm for A. peckiana; 10.0–12.0 × 5.5–7.0 μm, Qm = 1.81 ± 0.14 for A. pinophila) [2,53].
Amanita fulvisquamea Yuan S. Liu and S. Lumyong, sp. nov.; Figure 2b–d and Figure 4.
MycoBank number: 847955
Etymology: “fulvisquamea”, from fulvus (brownish) and squameus (covered with scales), referring to the brown scales on its pileus.
Holotype: THAILAND, Phetchabun Province, Nam Nao District, 16°42′37″ N 101°35′55″ E, alt. 870 m, 21 August 2020, Yuan S. Liu, STO-2020-367 (CMUB39993). GenBank accession numbers: OQ780689 (ITS), OQ780671 (nrLSU), OQ740051 (RPB2), OQ740069 (TEF1-α), and OQ740087 (TUB).
Basidiomata small- to medium-sized. Pileus 3.0–5.8 cm in diam., plano-convex to applanate, white (1A1) with pale orange (5A3) tone; volval remnants on pileus floccose to scaly, white (1A1), greyish orange (6B4–6) to brown (6C4–6), densely arranged over the disk; margin inconspicuously striate at first and becoming obviously so with age, appendiculate; context 3.5–7.0 mm wide, white (1A1), unchanging. Lamellae free, crowded, white (1A1), becoming greyish orange (5B3–5) to brown (6E5–8) upon drying; lamellulae mostly truncate. Stipe 6.0–7.5 cm long × 0.5–1.2 cm diam. (the length includes the basal bulb), subcylindric or slightly tapering upwards, with the apex slightly expanded, white (1A1), covered with floccose, white (1A1), greyish orange (6B4–6) to brown (6C4–6) squamules; context white (1A1), unchanging, fistulose; basal bulb absent; volva saccate, 2.5–3.1 cm high × 1.8–2.7 cm wide., membranous, white (1A1) to brown (6C4–6). Annulus present, white (1A1), fugacious. Odor not recorded.
Lamellar trama bilateral. Mediostratum 25–40 μm wide, consisting of abundant clavate to oblong inflated cells (35–145 × 12–35 μm); filamentous hyphae abundant, 2–7 μm wide; vascular hyphae scarce. Lateral stratum 35–55 μm thick, consisting of abundant oblong inflated cells (22–75 ×11–32 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–8 μm wide. Subhymenium 25–35 μm thick, with two–three layers of subglobose or irregular cells, 9–25 × 7–13 μm. Basidia (Figure 4b) 33–51 × 10–14 μm, clavate, four-spored; sterigmata 3–5 μm long; basal septa lacking clamps. Basidiospores [81/3/3] (8.0–) 8.5–11.0 (–11.5) × (6.5–) 7.0–8.0 (–8.5) μm, avl × avw = 9.7 × 7.2 μm, Q = (1.13–) 1.20–1.57 (–1.64), Qm = 1.35 ± 0.11, broadly ellipsoid to ellipsoid, sometimes subglobose or elongate, thin-walled, smooth, colorless to pale yellow or dull yellow, amyloid (Figure 4a). Lamellar edge sterile, consisting of subglobose to ellipsoid or pyriform inflated cells (18–65 × 11–30 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–5 μm wide, irregularly arranged. Pileipellis 110–170 μm thick, two-layered; upper layer (80–135 μm thick) strongly gelatinized, consisting of subradially arranged, thin-walled, colorless to pale yellow, filamentous hyphae 1–8 μm wide; lower layer (20–35 μm thick) consisting of radially and compactly arranged, filamentous hyphae 2–6 μm wide, colorless to pale yellow; vascular hyphae scarce. Volval remnants on pileus (Figure 4c) consisting of subradially to radially arranged elements: filamentous hyphae abundant, 2–8 μm wide, colorless to pale yellow, thin-walled; inflated cells abundant, ellipsoid to clavate, 43–200 × 13–37 μm, colorless to pale yellow, thin-walled, often terminal; vascular hyphae scarce. Interior of volval remnants on stipe base consisting of sub-longitudinally arranged elements: filamentous hyphae abundant, 2–12 μm wide, colorless, thin-walled; inflated cells abundant, subglobose to ellipsoid, 30–50 × 12–40 μm, colorless, thin-walled, often terminal; vascular hyphae scarce. The outer surface of volval remnants on stipe base consisting of very abundant filamentous hyphae (1–11 μm wide), mixed with scattered to fairly abundant, subglobose to ellipsoid, or ovoid to pyriform inflated cells. The inner surface of volval remnants on stipe base gelatinized, similar to structure of interior part but comprising much more filamentous hyphae (1–10 μm wide). Stipe trama consists of longitudinally arranged, long clavate terminal cells, 105–320 × 16–35 μm; filamentous hyphae scattered to abundant, 2–10 μm wide; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous or coniferous forests dominated by Dipterocarpus, Shorea, and Pinus species. Basidiomata occurs in the rainy season.
Distribution: Currently known in northern Thailand.
Additional collections examined: THAILAND, Chiang Mai Province, Mae On District, alt. 1201 m, 30 July 2020, Yuan S. Liu, STO-2020-211 (SDBR-STO20-211). Phetchabun Province, Nam Nao District, alt. 870 m, 21 August 2020, Yuan S. Liu, STO-2020-377 (SDBR-STO20-377).
Notes: Amanita fulvisquamea is characterized by its small- to medium-sized basidiomata, pulverulent to floccose, or patchy and greyish orange to brown volval remnants on pileus, striate pileal margin, saccate volva remnants on the stipe base, as well as broadly ellipsoid to ellipsoid basidiospores (8.5–11.0 × 7.0–8.0 μm, Qm = 1.35 ± 0.11).
Morphologically, Amanita fulvisquamea is easily confused with A. clarisquamosa, A. parvicurta, and A. volvata due to the similar appearances, e.g., floccose to pulverulent, brownish volval remnants on the pileus; floccose, brownish squamules on the stipe, as well as saccate volva remnants on the stipe base. However, A. clarisquamosa has much longer basidiospores, which mainly possess elongate shape, while both A. parvicurta and A. volvata have much narrower and elongate-shaped basidiospores [2,16,53].
Phylogenetically, Amanita brunneomaculata Y.Y. Cui, Q. Cai and Zhu L. Yang is closely related to A. fulvisquamea. However, the former differs from the latter by having a distinctly spotted pileus, as well as much longer basidiospores (10.0–13.0 × 6.5–8.0 μm, Qm = 1.65 ± 0.19) [2].
Amanita lanigera is morphologically similar and phylogenetically related to A. fulvisquamea. However, A. lanigera, originally reported from China, has thicker pileipellis (75–230 μm), larger inflated terminal cells (80–520 × 15–45 μm) in the stipe trama, as well as longer, ellipsoid, and colorless basidiospores (10.0–12.0 × 7.0–8.5 μm, Qm = 1.35 ± 0.11) [2].
Amanita sect. Phalloideae (Fr.) Quél., Mém. Soc. Emul. Montbéliard, Ser. II, 5: 66 (1872).
Basionym: Amanita [sect.] Phalloideae Fr., Monogr. Amanit. Sueciae: 3 (1854).
Lectotype: Amanita phalloides (Vaill. ex Fr.) Link., Handbuch zur Erkennung der Nutzbarsten und am häufigsten vorkommenden Gewächse: 272 (1833).
Notes: According to previous studies, Amanita sect. Phalloideae phylogenetically comprises three subclades that are well supported in phylogenetic analyses and by morphological evidence. Our multi-locus phylogenetic analysis also presented the same result. These three subclades may be treated as subsections or new sections [2,4,26,54] in the future.
Up to now, six taxa of section Phalloideae have been reported from Thailand, namely Amanita ballerina Raspé, Thongbai and K.D. Hyde, A. brunneitoxicaria Thongbai, Raspé and K.D. Hyde, A. fuliginea Hongo, A. fuligineoides P. Zhang and Zhu L. Yang, A. rimosa P. Zhang and Zhu L. Yang and A. zangii Zhu L. Yang, T.H. Li and X.L. Wu [25,26,29]. One more taxon is recognized in our phylogenetic analysis, and here it is described as a new species based on morphological evidence as well.
Amanita albifragilis Yuan S. Liu and S. Lumyong, sp. nov.; Figure 2e–f and Figure 5.
MycoBank number: 847956
Etymology: “albifragilis”, from albus (whitish) and fragilis (brittle), refers to the white fruiting body and the thin and brittle surface of the pileus.
Holotype: THAILAND, Sakon Nakhon Province, Kut Bak District, 17°6′4″ N 103°54′32″ E, alt. 205 m, 15 August 2020, Yuan S. Liu, STO-2020-300 (CMUB39994). GenBank accession numbers: OQ780692 (ITS), OQ780674 (nrLSU), and OQ740072 (TEF1-α).
Basidiomata small- to medium-sized. Pileus 3.2–5.2 cm in diam., plano-convex to applanate, often depressed at center, surface thin and fragile, white (1A1); volval remnants on pileus absent; margin non-striate, non-appendiculate; context 1.5–2.5 mm wide, white (1A1), unchanging. Lamellae free, crowded, white (1A1); lamellulae mostly truncate. Stipe 5.4–8.0 cm long × 0.5–0.6 cm diam. (the length includes the basal bulb), subcylindric or slightly tapering upwards, with apex slightly expanded, white (1A1), covered with fibrous, white (1A1), squamules; context white (1A1), unchanging, fistulose to solid; basal bulb subglobose; volva limbate, 1.3–1.8 cm high × 1.4–1.8 cm wide., membranous, white (1A1). Annulus subapical, membranous, white (1A1), persistent. Odor not recorded.
Lamellar trama bilateral. Mediostratum 20–30 μm wide, consists of abundant ellipsoid to elongate inflated cells (60–108 × 15–32 μm); filamentous hyphae abundant, 2–8 μm wide; vascular hyphae scarce. The lateral stratum consists of abundant ellipsoid to clavate inflated cells (30–60 × 11–22 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 2–6 μm wide. Subhymenium 20–30 μm thick, with two–three layers of subglobose to ellipsoid or irregular cells, 7–19 × 6–13 μm. Basidia (Figure 5b) 28–45 × 8–12 μm, clavate, four-spored; sterigmata 3–6 μm long; basal septa lacking clamps. Basidiospores [69/2/2] (7.0–) 8.0–9.0 (–10.5) × 6.0–7.0 (–9.0) μm, avl × avw = 8.5 × 6.8 μm, Q = (1.13–) 1.14–1.39 (–1.50), Qm = 1.26 ± 0.09, broadly ellipsoid, sometimes subglobose, thin-walled, smooth, colorless, and amyloid (Figure 5a). Lamellar edge sterile, consisting of subglobose to ellipsoid or clavate inflated cells (14–50 × 12–28 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae scattered, 1–3 μm wide, irregularly arranged. Pileipellis 60–110 μm thick, two-layered; upper layer (20–40 μm thick) slightly gelatinized, consisting of subradially arranged, thin-walled, colorless, filamentous hyphae 2–8 μm wide; lower layer (30–70 μm thick) consisting of radially and compactly arranged, filamentous hyphae 2–8 μm wide, colorless; vascular hyphae scarce. The interior of volval remnants on the stipe base consists of sub-longitudinally to irregularly arranged elements: filamentous hyphae very abundant, 3–11 μm wide, colorless, thin-walled; inflated cells scarce to scattered, clavate, 70–110 × 10–36 μm, colorless, thin-walled; vascular hyphae scarce. The outer surface of volval remnants on stipe base (Figure 5c) predominately consists of very abundant filamentous hyphae (3–12 μm wide), mixed with scarce, clavate inflated cells. The inner surface of volval remnants on the stipe base gelatinized, similar to structure of interior part but comprising much more filamentous hyphae (3–9 μm wide). Stipe trama consists of longitudinally arranged, abundant, long clavate terminal cells (80–285 × 12–23 μm); filamentous hyphae abundant to very abundant, 2–9 μm wide; vascular hyphae scarce. Annulus consists of radially arranged elements: inflated cells scattered, clavate, often terminal, 35–75 × 9–17 μm, colorless, thin-walled; filamentous hyphae very abundant to predominant, 2–9 μm wide, colorless, thin-walled; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: Currently known in northeastern Thailand.
Additional collections examined: THAILAND, Sakon Nakhon Province, Kut Bak District, alt. 205 m, 15 August 2020, Yuan S. Liu, STO-2020-304 (SDBR-STO20-304).
Notes: Amanita albifragilis is characterized by its small- to medium-sized basidiomata, slightly depressed pileal center, the thin and fragile surface of the pileus, a non-striate pileal margin, subglobose stipe base surrounded by limbate volva remnants, subapical and persistent annulus, as well as the broadly ellipsoid amyloid basidiospores (8.0–9.0 × 6.0–7.0 μm, Qm = 1.26 ± 0.09).
At first sight, Amanita rimosa, originally reported from China [55], resembles the newly described species rather strongly. Both species share a number of similar or identical features, e.g., a small and white basidioma, a smooth pileus with slightly rimose margin, the limbate volva remnants on the stipe base, and a membranous and persistent annulus [2,19,55]. However, A. rimosa has globose to subglobose basidiospores (7.0–8.5 × 6.5–8.0 μm, Qm = 1.08 ± 0.05) and a different structure of pileipellis containing more abundant ellipsoid to clavate inflated cells [2,19,55].
Except for Amanita rimosa, there are a number of taxa that have white basidiomata in section Phalloideae, e.g., A. exitialis Zhu L. Yang and T. H. Li, A. parviexitialis Q. Cai, Zhu L. Yang and Y.Y. Cui, A. virosa Bertillon and A. subjunquillea S. Imai. Among the above taxa, A. parviexitialis is easily confused with A. albifragilis due to its small basidioma, a smooth and depressed pileus, and the limbate volva remnants on the stipe base. However, A. parviexitialis usually has brownish tone in the pileal center, two-spored basidia, as well as much wider, subglobose basidiospores (7.5–9.5 × 7.0–9.0 μm, Qm = 1.09 ± 0.05) [2,19,56].
Phylogenetically, Amanita albifragilis is closely related to A. griseorosea Q. Cai, Zhu L. Yang and Y.Y. Cui and A. molliuscula Q. Cai, Zhu L. Yang and Y.Y. Cui. Amanita griseorosea can be easily distinguished from A. albifragilis by having a grayish-brown pileus with dark-gray fibrils and pinkish lamellae [2,19,38,56]. Compared to the newly described species, A. molliuscula has much more abundantly inflated cells in structures of pileipellis and in the interior of volval remnants on stipe base, as well as the wider and globose to subglobose basidiospores (7.5–9.0 × 7.0–8.0 μm, Qm = 1.07 ± 0.06) [2,38,56].
Amanita sect. Roanokenses Singer ex Singer, Sydowia 15: 67 (1962).
Synonym: Amanita subsect. Limbatulae Bas, Persoonia 5: 528 (1969).
Type: Amanita roanokensis Coker, J. Elisha Mitchell scient. Soc. 43: 141 (1927).
Notes: Amanita sect. Roanokenses is one of the most species-diverse sections in Amanita subgen. Amanitina. To date, nine species have been reported from Thailand, namely A. alboflavescens Hongo, A. atrobrunnea Thongbai, Raspé and K.D. Hyde, Amanita cf. oberwinkleriana, A. hongoi Bas, A. japonica Hongo ex Bas, A. macrocarpa W. Q. Deng, T. H. Li and Zhu L. Yang, A. manginiana sensu W.F. Chiu, A. pseudoporphyria Hongo and A. virgineoides Bas [2,25,26,28]. In this study, four taxa belonging to the section Roanokenses were recognized and are presented below.
Amanita caojizong Zhu L. Yang, Y.Y. Cui and Q. Cai, Fungal Divers. 91: 138 (2018). Figure 2g and Figure 6.
Basidiomata large. Pileus 9.5–12.0 cm diam., convex to plano-convex, milk white to greyish yellow (1B2–4) or greyish brown (5E3), possessing innate dark-grey radiating fibrils; volval remnants on pileus often absent; margin non-striate, appendiculate; context 8–9 mm wide, white (1A1), unchanging. Lamellae free, crowded, white (1A1); lamellulae attenuate. Stipe 14.3–20.0 cm long × 1.5–1.7 cm diam. The length includes the basal bulb, cylindrical or slightly tapering upwards with apex slightly expanded, white (1A1), covered with fibrous squamules; context solid, white (1A1); basal part 2.2–3.2 cm diam., fusiform to clavate; volval remnants on stipe base sheathed, membranous, with free limb up to 6.1 cm high, white (1A1). Annulus apical, white, fragile, and fugacious when mature. Odor not recorded.
Lamellar trama bilateral. Mediostratum 20–40 μm wide, consisting of abundant ellipsoid to elongate inflated cells (53–90 × 13–22 μm); filamentous hyphae abundant, 2–6 μm wide; vascular hyphae scarce. Lateral stratum 20–30 μm wide, consisting of abundant elongate to clavate inflated cells (36–65 × 12–22 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–5 μm wide. Subhymenium 20–30 μm thick, with two–three layers of subglobose, ovoid to ellipsoid, or irregular cells, 6–25 × 5–16 μm. Basidia (Figure 6b) 32–45 × 8–10 μm, clavate, four-spored; sterigmata up to 4–5 μm long; basal septa lacking clamps. Basidiospores [75/3/3] (6.0–) 6.5–8.0 (–9.0) × 5.0–7.0 μm, avl × avw = 7.5 × 6.0 μm, Q = (1.00–) 1.14–1.36 (–1.50) μm, Qm = 1.25 ± 0.10, broadly ellipsoid, sometimes globose to subglobose, thin-walled, smooth, colorless, amyloid (Figure 6a). Lamellar edge sterile, consisting of subglobose to ellipsoid inflated cells (9–17 × 8–14 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–7 μm wide, irregularly arranged. Pileipellis 90–150 μm thick, two-layered; upper layer (30–90 μm thick) strongly gelatinized, consisting of radially, thin-walled, colorless or light brownish, filamentous hyphae 2–5 μm wide; lower layer (45–110 μm thick) consisting of radially and compactly arranged filamentous hyphae 2–7 (–12) μm wide, yellowish to brownish; vascular hyphae scarce. The inner part of volval remnants on the stipe base consists of longitudinally arranged elements: filamentous hyphae predominant, 2–8 μm wide, colorless, thin-walled, branching; inflated cells scarce to scattered, ellipsoid to clavate, sometimes subglobose, 65–125 × 13–30 μm, colorless, thin-walled, interjacent, or terminal; vascular hyphae scarce. Th outer surface of volval remnants on the stipe base (Figure 6c) is similar to the inner part but with more abundant filamentous hyphae. The inner surface of volval remnants on the stipe base is similar to the interior part but slightly gelatinized. Stipe trama consists of longitudinally arranged, long clavate terminal cells, 130–220 × 15–30 μm; filamentous hyphae abundant, 2–10 μm wide; vascular hyphae scarce. Annulus consists of loosely arranged, interwoven elements: inflated cells abundant, globose, subglobose to pyriform, 15–55 × 13–48 μm, colorless, thin-walled; filamentous hyphae fairly abundant, 1–6 μm wide, colorless, thin-walled; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [2,57], Japan [58], Korea [59], and Thailand ([26], this study).
Specimens examined: THAILAND, Chiang Mai Province, Mae Taeng District, alt. 720 m, 9 August 2019, Yuan S. Liu, STO-2019-473 (SDBR-STO19-473); Mae On District, alt. 753 m, 6 July 2020, Yuan S. Liu, STO-2020-120 (SDBR-STO20-120). Chiang Rai Province, Mae Fa Luang District, alt. 1236 m, 10 July 2020, Yuan S. Liu, STO-2020-169 (SDBR-STO20-169).
Notes: Amanita caojizong, reported from China, is a common edible mushroom found in Yunnan province. It is morphologically similar to a number of taxa, such as A. pseudoporphyria, A. pseudomanginiana Q. Cai, Y.Y. Cui and Zhu L. Yang, A. griseoturcosa T. Oda, C. Tanaka and Tsuda, A. roseolifolia Y.Y. Cui, Q. Cai and Zhu L. Yang and A. modesta Corner and Bas. Detailed comparisons between A. caojizong and these similar species can be found in Cui et al. [2]. It is worth noting that our Thai collections had a much wider color range on the pileus, i.e., milk white to greyish yellow or greyish brown.
Amanita griseofarinosa Hongo, Mem. Fac. Lib. Arts Shiga Univ. Nat. Sci. 11: 39 (1961). Figure 2h and Figure 7.
Basidiomata small- to medium-sized. Pileus 3.5–6.5 cm diam., convex to plano-convex, or applanate to plano-concave, light grey (4C1–2) to brownish grey (4D1–2); volval remnants on pileus floccose to pulverulent, brownish grey (4D2–3), yellowish (5D3–5) to yellowish brown (5E4–5), densely arranged on the disc; margin non-striate, appendiculate; context 3–4.5 mm wide, white (1A1), unchanging. Lamellae free, crowded, white (1A1); lamellulae attenuate. Stipe 7.2–12.0 cm long × 0.5–1.2 cm diam. (the length includes the basal bulb), cylindrical, densely covered by floccose to pulverulent yellowish-white (4A2) to orange-white (5A2) squamules; context stuffed, white (1A1); basal part 0.9–1.8 cm diam., clavate to ventricose, upper part covered with floccose to pulverulent, yellowish-white (4A2) to orange-white (5A2) volval remnants. Annulus fragile and fugacious. Odor not recorded.
Lamellar trama bilateral. Mediostratum 25–35 μm wide, consisting of abundant ellipsoid inflated cells (45–95 × 10–22 μm); filamentous hyphae abundant, 2–7 (–11) μm wide; vascular hyphae scarce. Lateral stratum 20–30 μm wide, consisting of abundant clavate inflated cells (35–85 × 8–25 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–7 μm wide. Subhymenium 20–30 μm thick, with two–three layers of subglobose to ellipsoid or irregular cells, 6–20 × 6–12 μm. Basidia (Figure 7b) 40–56 × 11–13 μm, clavate, four-spored; sterigmata up to 3–5 μm long; basal septa lacking clamps. Basidiospores [68/2/2] (7.0–) 7.5–10.0 (–11.0) × (6.0–) 6.5–8.5 (–9.0) μm, avl × avw = 8.6 × 7.4 μm, Q = (1.00–)1.06–1.31 (–1.33) μm, Qm = 1.17 ± 0.09, mainly subglobose to broadly ellipsoid, sometimes globose or ellipsoid, thin-walled, smooth, colorless, amyloid (Figure 7a). Lamellar edge sterile and consists of subglobose to ellipsoid inflated cells (15–33 × 10–30 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–6 μm wide, irregularly arranged. Pileipellis 50–100 μm thick, consisting of radially and compactly, thin-walled, colorless to light brownish, filamentous hyphae 2–5 μm wide, with its outer-surface hyphae loosely and irregularly arranged; vascular hyphae scarce to scattered. Volval remnants on pileus (Figure 7c) composed of abundant irregularly arranged filamentous hyphae 2–7 μm wide, mixed with abundant to predominant subglobose, or broadly clavate to fusiform inflated cells (15–53 × 10–40 μm). Volval remnants on stipe base similar to structure of volval remnants on pileus, predominately composed of irregularly arranged subglobose, or broadly clavate to fusiform inflated cells (9–50 × 8–32 μm), mixed with abundant filamentous hyphae (2–7 μm wide). Stipe trama consists of abundant longitudinally arranged, long clavate terminal cells, 80–220 × 19–30 μm; filamentous hyphae abundant, 3–10 μm wide; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [2,18,19], Japan [16,60,61], Korea [62], and Thailand [this study].
Specimens examined: THAILAND, Chiang Mai Province, Mae Taeng District, alt. 1077 m, 31 May 2020, Yuan S. Liu, STO-2020-8 (SDBR-STO20-08); Yuan S. Liu, STO-2020-9 (SDBR-STO20-09).
Notes: Amanita griseofarinosa was first reported from Japan [60] and was then found in China and Korea [2,18,19,62]. Our two specimens possess small- to medium-sized basidiomata, pileus and stipe densely covered by floccose to pulverulent, brownish-grey or yellowish-white squamules, appendiculate margin, attenuate lamellulae, and fragile and fugacious annulus. All these features are consistent with the type specimen.
Morphologically, the species is similar to Amanita cf. griseofarinosa (HKAS 79587) and A. vestita Corner and Bas on account of brownish-gray basidiomata, and pulverulent volval remnants densely pervade the surface of the pileus and stipe [2,15,19]. However, Amanita cf. griseofarinosa differs from A. griseofarinosa in its original sense by having clamps, as well as the much wider and globose to subglobose basidiospores (8.5–10.5 × 8.5–10.0 μm, Qm = 1.04 ± 0.04) [2,19]. Moreover, Amanita cf. griseofarinosa is phylogenetically distinct from A. griseofarinosa [2]. Amanita vestita, reported from Singapore, is distinguished from A. griseofarinosa by having small basidiomata, slightly depressed pileal center, and much narrower basidiospores (7.5–9.0 × 5.5–6.5 μm) [15].
Undoubtedly, Amanita berkeleyi (Hook. f.) Bas, originally described from India, is closely related to A. griseofarinosa [16]. However, A. berkeleyi possesses large to very large basidiomata, felted-pulverulent to crust-like volva remnants on its pileus, and much wider basidiospores (8.0–10.5 × 6.5–9.5 μm) [16].
Amanita cinereovelata Hosen, reported from Bangladesh [63], is phylogenetically related and morphologically similar to A. griseofarinosa. However, A. cinereovelata differs by having a thicker pileipellis (up to 290 μm), globose to subglobose basidiospores (9.0–10.0 × 8.0–9.0 μm, Qm = 1.12 ± 0.05), and the presence of clamps [63].
Amanita neoovoidea Hongo, Mem. Shiga Univ. 25: 57 (1975). Figure 2i and Figure 8.
Basidiomata large. Pileus 9.0–10.0 cm diam., convex to plano-convex, white (1A1) to yellowish white (1A2); volval remnants on pileus consisting of two layers: outer layer membranous, yellowish white to pale yellow (1A2–3); inner layer floccose to pulverulent, white (1A1); margin non-striate, appendiculate; context 8.5–10.0 mm wide, white (1A1), unchanging. Lamellae free, crowded, white (1A1); lamellulae attenuate. Stipe 12.0–13.5 cm long × 1.5–1.6 cm diam. (the length includes the basal bulb), cylindrical, densely covered by floccose to pulverulent white (1A1) squamules; context solid, white (1A1); basal part 2.6–3.0 cm diam., fusiform to ventricose; volval remnants on stipe base yellowish white (3A2), arranged in incomplete belts or with a recurved friable limb. Annulus subapical, white (1A1), fragile and fugacious. Odor not recorded.
Lamellar trama bilateral. Mediostratum 25–40 μm wide, consisting of abundant clavate to fusiform inflated cells (35–110 × 12–20 μm); filamentous hyphae abundant, 2–6 μm wide; vascular hyphae scarce. Lateral stratum 20–30 μm wide, consisting of abundant elongate to clavate inflated cells (20–145 × 12–30 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–7 μm wide. Subhymenium 20–30 μm thick, with two–three layers of subglobose, ovoid to ellipsoid or irregular cells, 6–27 × 6–16 μm. Basidia (Figure 8b) 40–55 × 10–12 μm, clavate, four-spored; sterigmata up to 3–5 μm long; basal septa lacking clamps. Basidiospores [52/2/2] 7.0–10.0 × 5.0–7.0 μm, avl × avw = 8.5 × 6.1 μm, Q = (1.14–)1.21–1.67 (–1.70) μm, Qm = 1.41 ± 0.12, broadly ellipsoid, sometimes subglobose or elongate, thin-walled, smooth, colorless, amyloid (Figure 8a). Lamellar edge sterile, consisting of pyriform to subglobose or clavate inflated cells (27–50 × 18–36 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–5 μm wide, irregularly arranged. Pileipellis 45–120 μm thick, consisting of radial, strongly gelatinized, colorless filamentous hyphae (2–6 μm wide); vascular hyphae scarce. Volval remnants on pileus (Figure 8c) consist of two layers. The outer layer of volval remnants on the pileus consists of more or less radially arranged elements: inflated cells (45–173 × 15–32 μm) fairly abundant to abundant; filamentous hyphae (2–9 μm wide) very abundant; vascular hyphae scarce. The inner layer of volval remnants on the pileus consists of irregularly arranged elements: inflated cells (14–83 × 10–50 μm) abundant to predominant, subglobose to clavate; filamentous hyphae (3–8 μm wide) abundant; vascular hyphae scarce. Volval remnants on stipe base composed of irregularly to vertical-arranged elements: inflated cells (36–105 × 16–25 μm) fairly abundant to abundant; filamentous hyphae (1.5–7.0 μm wide) very abundant to dominate; vascular hyphae scarce. Stipe trama consists of longitudinally arranged, long clavate terminal cells, 180–240 × 13–25 μm; filamentous hyphae abundant, 3–10 μm wide; vascular hyphae scarce. Annulus consists of loosely, irregularly arranged elements: inflated cells abundant, subglobose, ellipsoid to clavate, 23–70 × 15–32 μm, colorless, thin-walled; filamentsous hyphae scarce to fairly abundant, 1.5–5.0 μm wide, colorless, thin-walled; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [2,17,18,19], Nepal [64], Japan [64,65], Korea [66], and Thailand (this study).
Specimens examined: THAILAND, Chiang Mai Province, Mae On District, alt. 704 m, 6 July 2020, Yuan S. Liu, STO-2020-110 (SDBR-STO20-110); Mae Taeng District, alt. 720 m, 9 August 2019, Yuan S. Liu, STO-2019-503 (SDBR-STO19-503).
Notes: Amanita neoovoidea, originally described from Japan [65], is characterized by medium-sized to large basidiomata, pileal volval remnants arranged in two layers, with the outer layer being membranous and inner layer being floccose to pulverulent, appendiculate pileal margin, stipe densely covered by floccose to pulverulent squamules, with incomplete belts or recurved friable limb remnants on the stipe base, and a fragile and fugacious annulus. Our Thai materials are consistent with all the features above.
Morphologically, Amanita duplex Corner and Bas, reported from Singapore, is undoubtedly similar to this species. Moreover, in our phylogenetic analysis, A. neoovoidea is related to A. pseudomanginiana, A. pseudoporphyria, and A. atrobrunnea. Detailed comparisons between A. neoovoidea and the four related species above can be found in previous studies [2,18].
Amanita oberwinkleriana Zhu L. Yang and Yoshim. Doi, Bull. Natn. Sci. Mus. Tokyo 25 (3): 120 (1999). Figure 2j and Figure 9.
Basidiomata small- to medium-sized. Pileus 4.5–6.0 cm diam., plano-convex to applanate, sometimes plano-concave, smooth, white (1A1), often yellowish white (1A2) to pale yellow (4A2–3) in the center; volval remnants on pileus often absent; margin non-striate, sometimes with inconspicuous stripes (ca. 0.2–0.3 R), non-appendiculate; context 2.0–4.0 mm wide, white (1A1), unchanging. Lamellae free, crowded, white to yellowish white (1A1–2); lamellulae attenuate. Stipe 8.2–9.5 cm long × 0.7–1.0 cm diam. (the length includes the basal bulb), cylindrical, covered by white (1A1), fibrous to tomentose squamules; context fistulose, white (1A1); basal part 1.6–2.0 cm diam., fusiform to napiform; volval remnants on stipe base limbate, membranous, with free limb up to ca. 1.5 cm high, both surfaces white (1A1). Annulus subapical, membranous, white (1A1). Odor not recorded.
Lamellar trama bilateral. Mediostratum 25–30 μm wide, consisting of abundant ellipsoid inflated cells (48–105 × 13–23 μm); filamentous hyphae abundant, 3–8 μm wide; vascular hyphae scarce. Lateral stratum 25–30 μm wide, consisting of abundant ellipsoid to clavate inflated cells (32–60 × 12–25 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–7 μm wide. Subhymenium 30–40 μm thick, with two–three layers of subglobose, ovoid to ellipsoid, or irregular cells, 10–20 × 7–17 μm. Basidia (Figure 9b) 35–44 × 9–11 μm, clavate, four-spored; sterigmata up to 4–6 μm long; basal septa lacking clamps. Basidiospores [52/2/2] (7.5–) 8.0–10.0 (–11.0) × 6.0–7.5 (–8.0) μm, avl × avw = 9.0 × 6.8 μm, Q = 1.20–1.50 μm, Qm = 1.33 ± 0.09, broadly ellipsoid to ellipsoid, thin-walled, smooth, colorless, amyloid (Figure 9a). Lamellar edge sterile, consisting of subglobose to ellipsoid inflated cells (12–37 × 9–25 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae fairly abundant, 3–5 μm wide, irregularly arranged. Pileipellis 65–90 μm thick, two-layered; upper layer (30–60 μm thick) strongly gelatinized, consisting of radial, thinwalled, colorless, filamentous hyphae 2–4 μm wide; lower layer (30–35 μm thick) consisting of radially and compactly arranged, filamentous hyphae 2–6 μm wide, colorless; vascular hyphae scarce. Volval remnants on pileus (Figure 9c) consisting of irregularly arranged elements: inflated cell (23–48 × 20–35 μm), fairly abundant, subglobose to ellipsoid, single or in chains of two–three, colorless; filamentous hyphae (3–10 μm wide) abundant to dominate, colorless or light yellow; vascular hyphae scarce. Inner part of volval remnants on stipe base consisting of longitudinally arranged elements: filamentous hyphae very abundant to predominant, 1–8 μm wide, colorless, thin-walled; inflated cells fairly abundant to abundant, subglobose to ovoid, or ellipsoid, 23–55 × 18–37 μm, colorless, thin-walled; vascular hyphae scarce. The outer surface of volval remnants on stipe base similar to structure of interior part but with more abundant inflated cells. Inner surface gelatinized, similar to structure of interior part but with a few inflated cells. Stipe trama consists of longitudinally arranged, long clavate terminal cells, 170–310 × 16–27 μm; filamentous hyphae abundant, 3–10 μm wide; vascular hyphae scarce. Annulus consisting of loosely arranged, interwoven elements: inflated cells abundant, pyriform to subglobose, 16–43 × 12–24 μm, single or in chains of two–three, colorless, thin-walled; filamentous hyphae scarce to fairly abundant, 2–8 μm wide, colorless, thin-walled; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [2,18,19,67], India [68], Japan [61], Korea [20], and Thailand ([26], this study).
Specimens examined: THAILAND, Chiang Mai Province, Mueang Chiang Mai District, alt. 1143 m, 3 August 2019, Yuan S. Liu, STO-2019-359 (SDBR-STO19-359); Yuan S. Liu, STO-2019-372 (SDBR-STO19-372).
Notes: Amanita oberwinkleriana was firstly reported from Japan [61] and has also been found in other Asian countries, e.g., China, India, Korea, and Thailand [2,18,19,20,26,67,68]. It is characterized by small- to medium-sized basidiomata, a smooth and white pileus often tinged yellowish in the center, non-appendiculate pileal margin, attenuate lamellulae, fusiform to napiform stipe base surrounded by limbate volval remnants, as well as the membranous annulus. Our Thai materials are consistent with all the above features.
Morphologically, Amanita oberwinkleriana can be easily confused with a number of species, having a white and smooth pileus, limbate volval remnants on the stipe base, and a membranous annulus, e.g., A. exitialis, A. rimosa, and A. virosa. However, A. exitialis distinctly differs from the newly described species by having two-spored basidia and much larger basidiospores (9.5–12.0 × 9.0–11.5 μm, Qm = 1.08 ± 0.04) [2,18,19,20,67]. Amanita rimosa can be distinguished from A. oberwinkleriana by having fissured pileal margin, as well as smaller and globose to subglobose basidiospores (7.0–8.5 × 6.5–8.0 μm, Qm = 1.08 ± 0.05) [2,19,55]. Amanita virosa differs by having obvious and concolorous squamules on its stipe, as well as much wider and globose to subglobose basidiospores (8.0–11.0 × 8.0–10.0 μm, Qm = 1.07 ± 0.05) [2,19].
Phylogenetically, Amanita oberwinkleriana is related to A. rubiginosa Q. Cai, Y.Y. Cui and Zhu L. Yang, A. avellaneifolia Zhu L. Yang, Y.Y. Cui and Q. Cai, and A. modesta. However, this species distinctly differs from the latter three taxa by its small- to medium-sized basidiomata, as well as a smooth and white pileus [2,15,19].
Amanita sect. Validae (Fr.) Quél., Mém. Soc. Emul. Montbéliard, Ser. II, 5: 69 (1872).
Basionym: Agaricus sect. Validae Fr., Monogr. Amanit. Sueciea: 10 (1854).
Lectotype: Amanita excelsa (Fr.) Bertill., Dictionnaire encyclopédique des sciences médicales 1 (3): 499 (1866).
Notes: Previously, six taxa belonging to Amanita sect. Validae have been reported from Thailand, namely A. castanea Thongbai, Tulloss, Raspé and K.D. Hyde, Amanita cf. spissacea S. Imai, A. flavipes S. Imai sensu lato, A. fritillaria (Berk.) Sacc., A. sculpta Corner and Bas, and A. sinocitrina Zhu L. Yang, Zuo H. Chen and Z.G. Zhang [25,26,29]. In this study, A. cacaina L.P. Tang, T. Huang and N.K. Zeng and A. citrinoannulata Y.Y. Cui, Q. Cai and Zhu L. Yang are recognized and reported as two new records in Thailand on the basis of the phylogenetic and morphological analyses.
Amanita cacaina L.P. Tang, T. Huang and N.K. Zeng, Frontiers Microbiol. 13: 3 (2023). Figure 2k and Figure 10.
Basidiomata is very large. Pileus 18.0–18.6 cm diam., plano-convex to applanate, brownish orange (7C3–6) to brown (7E5–7); volval remnants on pileus often pyramidal to verrucose, 2–9 mm high and 2–8 mm wide, sometimes scaly, yellowish white (4A2) to dark brown (7F7–8); margin non-striate, appendiculate; context 5–17.0 mm wide, yellowish white (4A2) to light brown (7D4–5). Lamellae free, crowded, reddish brown (8E7–8) to dark brown (8F7–8); lamellulae attenuate. Stipe 23.8–27.3 cm long × 2.0–2.5 cm diam. (the length includes the basal bulb), cylindrical, light brown (7D4–6), densely covered by floccose to pulverulent white (1A1) to reddish brown (8E7–8) squamules; context solid; basal part 5.3–5.6 cm diam., globose to subglobose, upper part covered with verrucose to squarish, reddish brown (8D5–6) warts. Annulus fragile and fugacious. Odor not recorded.
Lamellar trama bilateral. Mediostratum 20–35 μm wide, consisting of abundant ellipsoid or clavate to fusiform inflated cells (56–135 × 17–35 μm); filamentous hyphae abundant, 2–6 μm wide; vascular hyphae scarce. Lateral stratum 20–30 μm wide, consisting of abundant ellipsoid to fusiform inflated cells (48–73 × 13–23 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 3–6 μm wide. Subhymenium 25–40 μm thick, with two–three layers of subglobose, or irregular cells, 15–28 × 10–18 μm. Basidia (Figure 10b) 38–56 × 12–20 μm, clavate, four-spored; sterigmata up to 3–6 μm long; basal septa lacking clamps. Basidiospores [50/2/2] (8.0–) 8.5–10.5 (–11.0) × 8.0–10.0 μm, avl × avw = 9.2 × 8.9 μm, Q = 1.00–1.13 (–1.24) μm, Qm = 1.04 ± 0.05, globose to subglobose, sometimes broadly ellipsoid, thin-walled, smooth, colorless to pale yellow, amyloid (Figure 10a). Lamellar edge sterile, consisting of pyriform to subglobose, or ellipsoid to clavate inflated cells (13–41 × 9–32 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–6 μm wide, irregularly arranged. Pileipellis 70–150 μm thick, two-layered, consisting of radial filamentous hyphae (3–12 μm wide); vascular hyphae scarce. Volval remnants on pileus (Figure 10c) consisting of vertically arranged elements: inflated cell (26–150 × 22–87 μm), abundant to predominant, subglobose to ellipsoid, single or in chains of two–three, colorless or light brownish yellow; filamentous hyphae (3–9 μm wide) abundant, colorless or light yellow; vascular hyphae scarce. Volval remnants on stipe base predominately composed of irregularly arranged light-brownish yellow, pyriform to subglobose, or ellipsoid to fusiform inflated cells (17–72 × 13–67 μm), mixed with abundant filamentous hyphae (3–7 μm wide). Stipe trama consisting of longitudinally arranged, long clavate terminal cells, 170–330 × 18–40 μm; filamentous hyphae abundant, 2–9 (–14) μm wide; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous or coniferous forests dominated by Dipterocarpus, Shorea, and Pinus species, respectively. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [69], and Thailand [this study].
Specimens examined: THAILAND, Phetchabun Province, Nam Nao District, alt. 887 m, 17 August 2020, Yuan S. Liu, STO-2020-324 (SDBR-STO20-324); Yuan S. Liu, STO-2020-338 (SDBR-STO20-338).
Notes: Our Thai materials possess very large basidiomata, pyramidal to verrucose and dark-brown pileal remnants, non-striate and appendiculate margin, solid stipe densely covered with floccose to pulverulent white to reddish-brown squamules, globose to subglobose stipe base surrounded by verrucose to squarish, reddish-brown warts. All these features are consistent with the original description of Amanita cacaina, which was recently published in China in 2023 [69]. Morphologically, A. cacaina is similar to a number of taxa, e.g., A. pseudosculpta L.P. Tang and T. Huang, A. sculpta, and A. westii (Murrill) Murrill. Detailed comparisons between A. cacaina and the related species above can be found in Huang et al. [69].
Amanita citrinoannulata Y.Y. Cui, Q. Cai and Zhu L. Yang, Fungal Divers. 91: 186 (2018). Figure 2l and Figure 11.
Basidiomata is small. Pileus 2.4 cm diam., plano-convex, brownish orange (5C4–6) to yellowish brown (5E7–8); volval remnants on pileus floccose-felted, dark yellow (4C8), densely arranged over disk, often washed away by rain; margin non-striate, non-appendiculate; context white (8A1), slowly changing to pinkish white (8A2) after injury. Lamellae free, crowded, white (1A1); lamellulae attenuate. Stipe 4.2 cm long × 0.4 cm diam. (the length includes the basal bulb), subcylindrical or slightly tapering upwards, yellowish white (4A2), turning to pinkish white or pale red (8A2–3) when bruised, covered by yellowish-white (4A2) to brown (5E8), snakeskin-shaped squamules above the annulus and fibrous squamules under the annulus; context solid, slowly changing to pinkish white (8A2) after injury; basal part 0.8 cm diam., ventricose, upper part covered with irregular, floccose, yellowish-white (4A2) volval remnants. Annulus subapical, membranous, fugacious, yellowish white to pale yellow (3A2–3). Odor not recorded.
Lamellar trama bilateral. Mediostratum 20–30 μm wide, consisting of abundant ellipsoid or clavate inflated cells (40–85 × 14–28 μm); filamentous hyphae abundant, 2–6 μm wide; vascular hyphae scarce. Lateral stratum 25–40 μm wide, consisting of abundant ellipsoid to clavate or ovoid inflated cells (20–80 × 14–30 μm), diverging at an angle of about 45° to the mediostratum; filamentous hyphae abundant, 2–11 μm wide. Subhymenium 20–25 μm thick, with two–three layers of subglobose, or irregular cells, 7–23 × 6–15 μm. Basidia (Figure 11b) 30–40 × 8–12 μm, clavate, four-spored; sterigmata up to 3–4 μm long; basal septa lacking clamps. Basidiospores [51/1/1] (7.0–) 7.5–9.0 × 6.0–7.5 μm, avl × avw = 8.1 × 6.5 μm, Q = (1.07–)1.14–1.38 (–1.50) μm, Qm = 1.25 ± 0.09, subglobose to broadly ellipsoid, thin-walled, smooth, colorless to light brown, amyloid (Figure 11a). Lamellar edge sterile, consisting of globose to subglobose, or ellipsoid inflated cells (18–36 × 17–28 μm), single or in chains of two–three, thin-walled, colorless; filamentous hyphae abundant, 2–6 μm wide, irregularly arranged. Pileipellis 100–150 μm thick, two-layered; upper layer (40–70 μm thick) slightly gelatinized, consisting of subradially arranged, thin-walled, colorless to pale yellow, filamentous hyphae 2–10 μm wide; lower layer (60–80 μm thick) consisting of radially and compactly arranged, colorless to pale-yellow filamentous hyphae 2–11 μm wide; vascular hyphae fairly abundant. Volval remnants on pileus (Figure 11c) consisting of almost vertically arranged elements: inflated cell (17–52 × 15–40 μm), abundant to predominant, subglobose to globose, single or in chains of two–three, colorless or light brown; filamentous hyphae (2–6 μm wide) scattered, colorless or light yellow; vascular hyphae scarce. Volval remnants on stipe base similar to the remnants on pileus but with more filamentous hyphae. Stipe trama consists of longitudinally arranged, long clavate to fusiform terminal cells, 63–238 × 20–38 μm; filamentous hyphae abundant, 2–7 μm wide; vascular hyphae scarce. Annulus consisting of radially arranged elements: inflated cells scattered, subglobose to clavate or ellipsoid, 20–43 × 12–34 μm, colorless to light brown, thin-walled; filamentous hyphae very abundant to predominant, 2–6 μm wide, colorless, thin-walled; vascular hyphae scarce. Clamps absent in all parts of basidioma.
Habitat: Solitary to scattered on soil in tropical deciduous forests dominated by Dipterocarpus and Shorea species. Basidiomata occurs in the rainy season.
Distribution: This species is currently known in China [2], and Thailand [this study].
Specimens examined: THAILAND, Chiang Mai Province, Mae Taeng District, alt. 720 m, 9 August 2019, Yuan S. Liu, STO-2019-483 (SDBR-STO19-483).
Notes: Amanita citrinoannulata was reported from China [2]. Our Thai material has a small basidioma, brownish pileus covered with floccose-felted, with dark-yellow remnants; context of pileus changing to pinkish tone after injury; yellowish stipe always changing to pinkish white or pale red when bruised; snakeskin-shaped squamules above the annulus and fibrous squamules under the annulus. All these features are consistent with the type specimen.
Morphologically, Amanita citrinoannulata is similar to a number of taxa, e.g., A. flavorubens (Berk. and Mont.) Sacc., A. fritillaria Sacc. f. fritillaria, and A. spissacea. Detailed comparisons between A. citrinoannulata and the species mentioned above can be found in [2,19].
Phylogenetically, Amanita citrinoannulata is closely related to A. detersa Zhu L. Yang, Y.Y. Cui and Q. Cai, A. flavoconia G. F. Atk, and A. spissa (Fr.) P. Kumm. However, A. detersa, reported from China [2], has a grayish-toned pileus, yellowish volva remnants on both pileus and stipe base, and pileal context not changing color when injured. Amanita flavoconia, reported from America [70], possesses a pileus that is usually umbonate, the remnants on its pileus are absent or present as yellow patches or powder, and has much narrower basidiospores (6.0–9.0 × 4.0–6.0 μm) [70]. Amanita spissa differs in having a robust basidiomata, lacking a color change in the pileal context, the snakeskin-liked squamules on the stipe being absent, and a white annulus [2,19].

4. Discussion

In our multigene phylogenetic analysis, the species in each section of the Amanita subgen. Amanitina are clearly recognized, and the topological structure among these six sections is also clearly presented. In particular, the relationship of the sect. Phalloideae and the other three sections, viz. sect. Arenariae, sect. Strobiliformes, and sect. Validae, are clearly recognized. Meanwhile, the species in sect. Phalloideae are separated into three subclades, which is consistent with previous studies [2,4,26,54,71,72].
In addition, the Amanita ballerina–A. chuformis subclade and A. hesleri–A. zangii subclade are composed of non-lethal species, which differ morphologically from lethal species from the A. millsii–A. virosa subclade by having elongated to ventricose basal bulbs, striate and appendiculate margins, etc. [15,16,26,54]. Thus, it seems reasonable that two non-lethal subclades could be considered as one or two new sections [4,26]. On the other hand, the A. millsii–A. virosa subclade is composed of lethal species, which often contain several deadly Amanita cyclic peptides, e.g., α-amanitin, β-amanitin, phalloidin, and phallacidin [38,56,73,74,75]. Our two samples and one undescribed specimen (ECM3-PKA) isolated from Dipterocarpus alatus roots, representing A. albifragilis, nest into the A. millsii–A. virosa subclade and cluster together with two deadly species, viz. A. griseorosea and A. molliuscula. According to previous studies [38,56], A. griseorosea contains β-amanitin, and A. molliuscula contains both α-amanitin and β-amanitin. Thus, A. albifragilis probably contains both compounds or at least one of them, and this speculation will be clarified in our next project.

Author Contributions

Conceptualization, Y.S.L. and S.L.; methodology, Y.S.L.; formal analysis, Y.S.L.; resources, Y.S.L. and J.-K.L.; data curation, Y.S.L.; writing—original draft preparation, Y.S.L.; writing—review and editing, J.-K.L., J.K., N.S. and S.L.; supervision, S.L.; project administration, S.L.; funding acquisition, S.L., J.K. and N.S. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by grants from Chiang Mai University and partially supported by grants from Plant Genetic Conservation Project under the Royal initiative of Her Royal Highness Princess Maha Chakri Sirindhorn, Chiang Mai University (R000032658), and National Research Council of Thailand (N42A650198).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The DNA sequence data obtained from this study were deposited in GenBank under accession numbers: ITS (OQ780686–OQ780697); nrLSU (OQ780668–OQ780685); RPB2 (OQ740048–OQ740065); TEF1-α (OQ740066–OQ740084); TUB (OQ740085–OQ740093).

Acknowledgments

We are grateful to staffs of Chiang Mai University Haripunchai Campus for their excellent field assistance. We would like to thank the TA & RA Scholarship, graduate school, Chiang Mai University, for supporting our samples’ collection. Jean Evans I. Codjia and Soumitra Paloi are thanked for their valuable suggestions.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. RAxML tree based on a concatenated dataset (nrLSU + ITS + RPB2 + TEF1-a + TUB). Bootstrap values (BS) for ML ≥ 60% and posterior probabilities (PPs) for BI ≥ 0.95 are placed above or below the branches. Newly generated sequences are indicated in red, and sequences from type material are marked with (T). The tree is rooted with Amanita flavofloccosa (HKAS101443 and HKAS90174) and A. vittadinii (HKAS101430).
Figure 1. RAxML tree based on a concatenated dataset (nrLSU + ITS + RPB2 + TEF1-a + TUB). Bootstrap values (BS) for ML ≥ 60% and posterior probabilities (PPs) for BI ≥ 0.95 are placed above or below the branches. Newly generated sequences are indicated in red, and sequences from type material are marked with (T). The tree is rooted with Amanita flavofloccosa (HKAS101443 and HKAS90174) and A. vittadinii (HKAS101430).
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Figure 2. Fresh basidiomata of studied Amanita species. (a) A. claristriata (CMUB39992, holotype). (b–d) A. fulvisquamea [(b,c) CMUB39993, holotype; (d) SDBR-STO20-211)]. (e,f) A. albifragilis (CMUB39994, holotype). (g) A. caojizong (SDBR-STO20-120). (h) A. griseofarinosa (SDBR-STO20-08). (i) A. neoovoidea (SDBR-STO19-503). (j) A. oberwinkleriana (SDBR-STO19-359). (k) A. citrinoannulata (SDBR-STO19-483). (l) A. cacaina (SDBR-STO20-324). Scale bars: (a–k) = 3 cm, (l) = 9 cm.
Figure 2. Fresh basidiomata of studied Amanita species. (a) A. claristriata (CMUB39992, holotype). (b–d) A. fulvisquamea [(b,c) CMUB39993, holotype; (d) SDBR-STO20-211)]. (e,f) A. albifragilis (CMUB39994, holotype). (g) A. caojizong (SDBR-STO20-120). (h) A. griseofarinosa (SDBR-STO20-08). (i) A. neoovoidea (SDBR-STO19-503). (j) A. oberwinkleriana (SDBR-STO19-359). (k) A. citrinoannulata (SDBR-STO19-483). (l) A. cacaina (SDBR-STO20-324). Scale bars: (a–k) = 3 cm, (l) = 9 cm.
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Figure 3. Amanita claristriata (CMUB39992, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 3. Amanita claristriata (CMUB39992, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 4. Amanita fulvisquamea (CMUB39993, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 4. Amanita fulvisquamea (CMUB39993, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 5. Amanita albifragilis (CMUB39994, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of outer surface of volval remnants on stipe base. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 5. Amanita albifragilis (CMUB39994, holotype). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of outer surface of volval remnants on stipe base. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 6. Amanita caojizong (SDBR-STO20-120). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of outer surface of volval remnants on stipe base. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 6. Amanita caojizong (SDBR-STO20-120). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of outer surface of volval remnants on stipe base. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 7. Amanita griseofarinosa (SDBR-STO20-09). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 7. Amanita griseofarinosa (SDBR-STO20-09). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 8. Amanita neoovoidea (SDBR-STO19-503). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 8. Amanita neoovoidea (SDBR-STO19-503). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 9. Amanita oberwinkleriana (SDBR-STO19-372). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 9. Amanita oberwinkleriana (SDBR-STO19-372). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 10. Amanita cacaina (SDBR-STO20-324). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 10. Amanita cacaina (SDBR-STO20-324). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Figure 11. Amanita citrinoannulata (SDBR-STO19-483). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
Figure 11. Amanita citrinoannulata (SDBR-STO19-483). (a) Basidiospores. (b) Hymenium and subhymenium. (c) Longitudinal section of volval remnants on pileus. Scale bars: (a,b) = 10 μm, (c) = 50 μm.
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Table
Table 1. Species names, voucher numbers, countries, and GenBank accession numbers of the taxa used in this study.
Table 1. Species names, voucher numbers, countries, and GenBank accession numbers of the taxa used in this study.
Species NamesVoucher NumbersCountriesGenBank Accession Numbers
nrLSUITSRPB2TEF1-αTUB
Section Amidella
A. avellaneosquamosaHKAS77340ChinaKJ466483KJ466418KJ466648KJ481982KJ466562
A. avellaneosquamosaHKAS100602ChinaMH486379MH508258MH485873MH508681—
A. brunneomaculataHKAS68393ChinaMH486410MH508278MH485892MH508698MH485428
A. brunneomaculataHKAS70032 TChinaMH486411MH508279MH485893MH508699—
A. clarisquamosaHKAS29514ChinaAF024448————
A. claristriataCMUB39992 TThailandOQ780668OQ780686OQ740048OQ740066—
A. claristriataSDBR-STO20-407ThailandOQ780669OQ780687OQ740049OQ740067OQ740085
A. claristriataSDBR-STO20-408ThailandOQ780670OQ780688OQ740050OQ740068OQ740086
A. curtipesAH 31718SpainEF653961AY486233———
A. curtipesAH 31924SpainEF653960EF653963———
A. fulvisquameaCMUB39993 TThailandOQ780671OQ780689OQ740051OQ740069OQ740087
A. fulvisquameaSDBR-STO20-211ThailandOQ780672OQ780690OQ740052OQ740070—
A. fulvisquameaSDBR-STO20-377ThailandOQ780673OQ780691OQ740053OQ740071—
A. lanigeraHKAS89030 TChinaMH486621MH508420MH486074MH508880—
A. lanigeraHKAS97561ChinaMH486622MH508421MH486075—MH485591
A. peckianaRV5Aug96USAAF097387————
A. pinophilaHKAS70167 TChinaMH486759MH508504MH486178—MH485682
A. pinophilaHKAS71662ChinaMH486760MH508505MH486179—MH485683
A. ponderosaAH 19752SpainEF653958AY486234———
A. ponderosaAH 19699SpainEF653959EF653962———
A. volvataS. Harsch 304USAAF024485————
A. whetstoneaeRET 386-9USAKX061533KX061519———
A. whetstoneaeRET 697-1USAKX061531KX061518———
Section Arenariae
A. arenariaVPI 679 TAustraliaGQ925382GQ925393———
A. arenariaPERTH 09316213AustraliaMW793397MW795714MW820674MW820649MW820671
A. arenarioidesPERTH 07627025 TAustraliaMW775283MW775309MW820678MW820655MW820664
A. peltigeraPERTH 08793514AustraliaMN900625MN894307MN912054MN909824—
A. peltigeraPERTH 08793581AustraliaMN900627MN894321MN912056MN909826MN905762
A. pseudoarenariaPERTH 08606765 TAustraliaMW775284MW775312MW820681MW820656MW820669
A. sabulosaPERTH 09178759 TAustraliaMW775279MW775291MW820675MW820650MW820660
A. sabulosaPERTH 09178775AustraliaMW775280MW775296MW820677MW820653MW820661
Section Phalloideae
A. albifragilisCMUB39994 TThailandOQ780674OQ780692—OQ740072—
A. albifragilisSDBR-STO20-304ThailandOQ780675OQ780693—OQ740073OQ740088
A. albolimbataJEIC0667BeninMT966939MT966932MT966958MT966953MT966947
A. albolimbataJEIC0739 TBeninMT966942MT966935MT966963MT966955MT966950
A. ballerinaOR1026 TThailandNG_058607KY747467KY656884—KY656865
A. ballerinoidesJEIC0479GuineaOK510856OK510854OK510847OK510824OK510835
A. bweyeyensisJD 1304 TRwandaMK570927MK570920MK570937MK570940MK570916
A. bweyeyensisTS 591TanzaniaMK570928MK570921———
A. chuformisHKAS94075ChinaMT395380MT395378—MT364256—
A. chuformisHKAS101028 TChinaMT395381MT395379MT364258MT364257—
A. exitialisHKAS75774ChinaJX998052JX998027KJ466591JX998001KJ466503
A. exitialisHKAS75776ChinaJX998051JX998025KJ466593JX998003KJ466505
A. griseoroseaHKAS77333ChinaKJ466475KJ466412KJ466660—KJ466579
A. griseoroseaHKAS77334ChinaKJ466476KJ466413KJ466661—KJ466580
A. harkonenianaTS 1061 TTanzaniaMK570930MK570923———
A. hesleriRET 802-4USAMH836560MH836568———
A. levistriataFLAS-F-61208USAMH620278MH211813———
A. millsiiHO 581533 TAustraliaKY977713KY977714MF000753MF000759MF000760
A. millsiiHKAS 77322AustraliaKJ466457KJ466395KJ466643KJ481978KJ466557
A. molliusculaHKAS77324 TChinaKJ466472KJ466409KJ466639KJ481974KJ466553
A. molliusculaHMJAU20469ChinaKJ466473KJ466410KJ466640KJ481975KJ466554
A. rimosaHKAS75779ChinaJX998046JX998020KJ466617JX998004KJ466528
A. rimosaHKAS77336ChinaKJ466456KJ466394KJ466622KJ481958KJ466533
Amanita sp.ECM3-PKAThailand—DQ146367———
A. subpallidoroseaLHJ140923-41 TChinaKP691692KP691683KP691701KP691670KP691711
A. virosaHKAS90176ChinaMH486948MH508650MH486341MH509167MH485847
A. virosaHMJAU23304ChinaKJ466498KJ466431KJ466667KJ481999KJ466587
A. zangiiHKAS99663ChinaMH486958MH508655MH486351MH509178MH485855
A. zangiiHKAS77331ChinaKJ466500KJ466433KJ466669KJ482001KJ466589
Section Roanokenses
A. atrobrunneaBZ-N09ThailandKT934314KY747455KY656871—KY656852
A. avellaneifoliaHKAS80011 TChinaMH486378—MH485872MH508680MH485410
A. caojizongHKAS56933ChinaKJ466438KJ466378KJ466603KJ481943KJ466515
A. caojizongHKAS79673 TChinaMH486429MH508291MH485909MH508714—
A. caojizongSDBR-STO19-473ThailandOQ780676—OQ740054OQ740074OQ740089
A. caojizongSDBR-STO20-120ThailandOQ780677—OQ740055OQ740075OQ740090
A. caojizongSDBR-STO20-169ThailandOQ780678—OQ740056OQ740076—
A. cf. manginianaBZ-N11ThailandKY747474KY747457KY656873—KY656854
A. cf. oberwinklerianaBZ2013-39ThailandKY747476KY747459KY656876—KY656857
A. cinereovelataHKAS81647 TBangladeshKP259291—KP259288KP259289—
A. cinereovelataHKAS 81407BangladeshKP259292——KP259290—
A. eijiiHKAS70229ChinaMH486484MH508333MH485963MH508761MH485486
A. eijiiHKAS81225ChinaMH486485—MH485964MH508762MH485487
A. griseofarinosaHKAS80926ChinaMH486559MH508375MH486025MH508830MH485545
A. griseofarinosaHKAS81840ChinaMH486560—MH486026MH508831MH485546
A. griseofarinosaSDBR-STO20-08ThailandOQ780679—OQ740057OQ740077OQ740091
A. griseofarinosaSDBR-STO20-09ThailandOQ780680—OQ740058OQ740078OQ740092
A. kotohiraensisHKAS100500ChinaMH486613MH508414———
A. kotohiraensisHKAS101421ChinaMH486615—MH486069MH508875MH485587
A. manginianaHKAS101404ChinaMH486637————
A. manginianaMHHNU 30818ChinaMH605436—MH614263MH614264MH614265
A. modestaHKAS100505ChinaMH486647MH508438MH486098MH508904MH485613
A. neoovoideaHKAS89025ChinaMH486656MH508445MH486106MH508913MH485621
A. neoovoideaHKAS89065ChinaMH486657—MH486107MH508914MH485622
A. neoovoideaSDBR-STO19-503Thailand——OQ740060OQ740080—
A. oberwinklerianaHKAS85832ChinaMH486681—MH486118—MH485632
A. oberwinklerianaSDBR-STO19-359ThailandOQ780682OQ780694OQ740061OQ740081—
A. oberwinklerianaSDBR-STO19-372ThailandOQ780683OQ780695OQ740062OQ740082—
A. pseudomanginianaHKAS83470 TChinaMH486772—MH486187—MH485694
A. pseudoporphyriaBZ-N10ThailandKY747473KY747456KY656872—KY656853
A. pseudoporphyriaHKAS56984ChinaKJ466450KC429050KJ466614KJ481953KJ466525
A. rubiginosaHKAS52216 TChinaMH486817MH508561MH486229—MH485734
A. rubiginosaHKAS57045ChinaMH486819MH508563MH486231—MH485736
A. timidaHKAS83228ChinaMH486930MH508636MH486323MH509147MH485830
A. timidaHKAS89001ChinaMH486932—MH486325MH509149—
Section Strobiliformes
A. aspericepsHKAS80014ChinaMH486374MH508257MH485868MH508676MH485408
A. cinereoradicataHKAS101435ChinaMH486451MH508307MH485933MH508729MH485458
A. cinereoradicataHKAS69389ChinaMH486453MH508308MH485934MH508730MH485459
A. griseoverrucosaHKAS57357ChinaMH486581MH508392MH486043MH508850MH485562
A. strobiliformisHKAS84872GermanyMH486895MH508614MH486298MH509117MH485798
Section Validae
A. cacainaMHKMU NK Zeng 2557ChinaON768725ON768705———
A. cacainaSDBR-STO20-324ThailandOQ780684OQ780696OQ740063OQ740083—
A. castaneaMFLU 15-1424ThailandKU877539KU904823———
A. citrinaHKAS53467GermanyMH486457MH508312MH485937MH508733MH485461
A. citrinaHKAS101397FranceMH486456MH508311MH485936MH508732MH485460
A. citrinoannulataHKAS81994ChinaMH486463MH508317MH485943MH508739MH485466
A. citrinoannulataHKAS100524ChinaMH486459MH508314MH485939MH508735MH485463
A. citrinoannulataSDBR-STO19-483ThailandOQ780685OQ780697OQ740065OQ740084—
A. detersaHKAS71476 TChinaMH486475MH508328MH485954MH508752MH485479
A. detersaHKAS83720ChinaMH486479MH508332MH485958MH508756MH485482
A. flavoconiaRET 439-8USAMH486511MH508348MH485983MH508787—
A. flavoconiaRET 450-10USAMH486512MH508349MH485984MH508788—
A. sepiaceaHKAS56799ChinaMH486847MH508584MH486256—MH485759
A. sepiaceaHKAS68614ChinaMH486851MH508585MH486260—MH485763
A. spissaHKAS92051ChinaMH486892MH508611MH486295MH509114MH485795
Outgroup
A. flavofloccosaHKAS90174ChinaKT833801MH508352KT833818KT833831MH485508
A. flavofloccosaHKAS101443ChinaMH486515—MH485986MH508791MH485507
A. vittadiniiHKAS101430ItalyMH486950MH508651MH486342MH509169—
Newly generated sequences in this study are in red. Holotypes are marked with “T”.
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MDPI and ACS Style

Liu, Y.S.; Liu, J.-K.; Kumla, J.; Suwannarach, N.; Lumyong, S. Taxonomic Novelties and New Records of Amanita Subgenus Amanitina from Thailand. J. Fungi 2023, 9, 601. https://doi.org/10.3390/jof9060601

AMA Style

Liu YS, Liu J-K, Kumla J, Suwannarach N, Lumyong S. Taxonomic Novelties and New Records of Amanita Subgenus Amanitina from Thailand. Journal of Fungi. 2023; 9(6):601. https://doi.org/10.3390/jof9060601

Chicago/Turabian Style

Liu, Yuan S., Jian-Kui Liu, Jaturong Kumla, Nakarin Suwannarach, and Saisamorn Lumyong. 2023. "Taxonomic Novelties and New Records of Amanita Subgenus Amanitina from Thailand" Journal of Fungi 9, no. 6: 601. https://doi.org/10.3390/jof9060601

APA Style

Liu, Y. S., Liu, J. -K., Kumla, J., Suwannarach, N., & Lumyong, S. (2023). Taxonomic Novelties and New Records of Amanita Subgenus Amanitina from Thailand. Journal of Fungi, 9(6), 601. https://doi.org/10.3390/jof9060601

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