Corresponding author: Yupeng Ge (
Academic editor: Thorsten Lumbsch
Bluish
Na Q, Liu Z, Zeng H, Ke B, Song Z, Cheng X, Ge Y (2022) Taxonomic studies of bluish
Eight bluish
To date, fewer than 100 species of
Macromorphological observations were made on fresh specimens in the field and from photographs, with colour terms and notation following
Genomic DNAs of the putative new species were extracted from dried materials using a NuClean PlantGen DNA kit (Kangwei Century Biotechnology, Beijing, China). The internal transcribed spacer (
Specimens along with GenBank accession numbers used in the phylogenetic analysis. Sequences newly generated in this study are indicated in bold.
No. | Species | Voucher | Origin | LSU ID | References | |
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1. |
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231a | Venice |
|
— | Unpublished |
2. |
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HMJAU 43282 | China |
|
|
Unpublished |
3. |
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HMJAU 43468 | China |
|
— | Unpublished |
4. |
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KA12-0434 | Korea |
|
— |
|
5. |
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Aronsen120803 | Norway |
|
— |
|
6. |
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Orstadius329-05 | Norway |
|
— |
|
7. |
|
Aronsen061119 | Norway |
|
— |
|
8. |
|
Aronsen120826 | Norway |
|
— |
|
9. |
|
MGW1504 | USA |
|
|
Unpublished |
10. |
|
SAT1518708 | USA |
|
|
Unpublished |
11. |
|
CM14-RG2 | USA |
|
— | Unpublished |
12. |
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189f | Italy |
|
— |
|
13. |
|
4745-HRL 1312 | Canada |
|
— | Unpublished |
14. |
|
CBS 352.50 | France |
|
— |
|
15. |
|
CBS 254.53 | France |
|
— |
|
16. |
|
CBS 257.53 | France |
|
|
|
17. |
|
H6036851 | Finland |
|
— | Unpublished |
18. |
|
252b | Italy |
|
— |
|
19. |
|
252f | Italy |
|
— |
|
20. |
|
|
|
|
|
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21. |
|
|
|
|
|
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22. |
|
|
|
|
|
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23. |
|
|
|
|
|
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24. |
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ACL257 | Malaysia |
|
— |
|
25. |
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ACL271 | Malaysia |
|
— |
|
26. |
|
Aronsen051014 | Norway |
|
— |
|
27. |
|
173 | Russia |
|
— |
|
28. |
|
317h | Italy |
|
— |
|
29. |
|
AD4TN | Tunisia |
|
— | Unpublished |
30. |
|
615i | Italy |
|
— |
|
31. |
|
615b | Italy |
|
— |
|
32. |
|
120b | Italy |
|
— |
|
33. |
|
DM334g | India |
|
— |
|
34. |
|
KA13-1230 | Korea |
|
— |
|
35. |
|
320f | Italy |
|
— |
|
36. |
|
HMJAU 43048 | China |
|
— |
|
37. |
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HMJAU 43052 | China |
|
|
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38. |
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HMJAU 43126 | China |
|
|
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39. |
|
3782 | Canada |
|
— | Unpublished |
40. |
|
KA12-1699 | Korea |
|
— |
|
41. |
|
287f | Italy |
|
— |
|
42. |
|
DM136-40516 | USA |
|
— | Unpublished |
43. |
|
LXL71 | China |
|
— | Unpublished |
44. |
|
F26441 | USA |
|
— | Unpublished |
45. |
|
EP.19-A1625 | Greece |
|
— | Unpublished |
46. |
|
50 | Norway |
|
— | Unpublished |
47. |
|
TFB14649 | USA |
|
— | Unpublished |
48. |
|
ACL161 | Malaysia |
|
— |
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49. |
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ACL175 | Malaysia |
|
— |
|
50. |
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ACL212 | Malaysia |
|
— |
|
51. |
|
1028 | Italy |
|
— |
|
52. |
|
CNH03 ( |
USA |
|
— | Unpublished |
53. |
|
39 | Italy |
|
— |
|
54. |
|
39d | Italy |
|
— |
|
55. |
|
313b | Italy |
|
— |
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56. |
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GG436-86 | Svalbard |
|
— |
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57. |
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CBS 228.47 | France |
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|
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58. |
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CBS 229.47 | France |
|
|
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59. |
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HK47-15 | Norway |
|
— |
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60. |
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979a | Italy |
|
— |
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61. |
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FCME25817 | USA |
|
— |
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62. |
|
USA |
|
— |
|
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63. |
|
USA |
|
— |
|
|
64. |
|
CBH164 | Denmark |
|
— | Unpublished |
65. |
|
108b | Italy |
|
— |
|
66. |
|
108f | Italy |
|
— |
|
67. |
|
JN198391 | China |
|
— |
|
68. |
|
420526MF0010 | China |
|
— | Unpublished |
69. |
|
439b | Italy |
|
— |
|
70. |
|
439f | Italy |
|
— |
|
71. |
|
USA |
|
— |
|
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72. |
|
CBH410 | USA |
|
— | Unpublished |
73. |
|
SL09-06 | Canada |
|
— | Unpublished |
74. |
|
G. Alfredsen | Norway |
|
— | Unpublished |
75. |
|
938a | Italy |
|
— |
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76. |
|
Champ-21 | Spain |
|
— |
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77. |
|
407q | Italy |
|
— |
|
78. |
|
TL-12780 | USA |
|
|
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79. |
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ACL136 | Malaysia |
|
|
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80. |
|
ACL308 | Malaysia |
|
|
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81. |
|
71l | Italy |
|
— |
|
82. |
|
71h | Italy |
|
— |
|
83. |
|
515 | Italy |
|
— |
|
84. |
|
CC 13-12 | USA |
|
— |
|
85. |
|
455 | Italy |
|
— |
|
86. |
|
|
|
|
|
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87. |
|
|
|
|
|
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88. |
|
|
|
— |
|
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89. |
|
128a | Italy |
|
— |
|
90. |
|
p187i | USA |
|
— | Unpublished |
91. |
|
OSC 113746 | USA |
|
— | Unpublished |
92. |
|
104h | Italy |
|
— |
|
93. |
|
447h | Italy |
|
— |
|
94. |
|
3781 | Canada |
|
— | Unpublished |
95. |
|
KA13-1265 | Korea |
|
— |
|
96. |
|
TFB14487 | USA |
|
|
|
97. |
|
TFB7283A | USA |
|
|
|
Phylogenetic tree inferred from partial
In the tree shown in Fig.
It is noteworthy that the six samples of
In addition to morphological studies of the new taxa collected in China, morphological observations were made on 17 bluish specimens of
Our morphological observations using a light microscope confirmed the identity of 12 specimens as
This species is characterised by blue pileus, turning bluish-grey with age, pileus covered by a separable, gelatinous pellicle, stipe pruinose and with a blue base and stipe basal disc and acanathocysts of pileipellis absent.
China. Ningxia Hui Autonomous Region: Liangdianxia, Liupan Mountains National Forest Park, Jingyuan County, Guyuan City,
Refers to the pileus colour: blue when young, becoming bluish-grey with age.
Pileus 12–25 mm in diameter, hemispherical when young, conical, obtusely conical, campanulate with age, shallowly sulcate, translucently striate, almost smooth when young, becoming slightly brownish scaly at the centre, pruinose, with a glabrescent margin, dull blue (23D5) at the centre, margin pallescent to pastel blue (23A4), turning bluish-grey (23D2–23D3), a bit sticky, covered by a separable, gelatinous pellicle. Context white, thin, fragile. Lamellae 16–28 reaching the stem, adnate to slightly adnexed with a short tooth, narrowly spaced, white, with intervenose veins, edges concolorous with the face. Stipe 48–76 × 1.5–2.0 mm, equal or slightly broadened below, hollow, fragile, entirely pruinose (Fig.
Fresh basidiomata of
Basidiospores [60/3/2] (8.8) 9.3–
Microscopic features of
Scattered on humus and fallen leaves in mixed forests of
Morphological features of
Ningxia Hui Autonomous Region, China.
Ningxia Hui Autonomous Region: Xiaonanchuan, Jingyuan County, Guyuan City, 20 July 2020, Qin Na, Yupeng Ge, Hui Zeng, Junqing Yan and Zewei Liu,
The original description of
Morphological comparison of
Taxa |
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12–25 mm diam., hemispherical when young, conical, obtusely conical, campanulate with age, smooth when young, becoming slightly brownish scaly at the center, pruinose, acid blue to dull blue at the center and margin pallescent, turning bluish gray, covered by a separable, gelatinous pellicle. | 3.5–13 mm in diam., parabolic, obtusely conical when young, hemispherical, campanulate with age, with an umbo at the center, shallowly sulcate, translucent-striate, smooth, gelatinous slightly, the margin infrequently out of flatness, brown to dark brown, becoming acid blue to dull blue towards the margin, with a greyish white margin, covered by a separable, sticky pellicle. | (3)5–15(25) mm broad, more or less ovoid with an appressed or slightly incurved margin, becoming obtusely conic to campanulate, surface lubricous subviscid, glabrous or appearing somewhat granulose near the margin, translucent-striate, pellicle tenacious and completely separable, pale blue or greenish blue, soon tinged with brown and assuming various degrees of bluish, greenish, or grayish brown with a pallid margin, often sordid yellowish in age, bluish tints often lingering on the margin. | 5–15 mm wide, conical to campanulate, ± sulcate, translucent-striate, finely puberulous, covered with a separable gelatinous pellicle, pale grey-brown or pale sepia brown, sometimes with an olivaceous, greenish or bluish green shade, margin often bluish green, or more rarely dingy citrine to ochraceous yellow. | 2–5(–10) mm wide, covered with a (separable), gelatinous pellicle, at first ± globose, then hemispherical to parabolic, becoming convex or somewhat depressed, but also with a small papilla, sulcate, translucent-striate, pruinose, glabrescent, somewhat lubricous, initially pale brown, then pale grey with darked centre, becoming almost white with age. | 16 mm in diam., up to 4 mm high, at first subglobose to ovoid-conical, with age becoming convex to shallowly so, slightly depressed at apex, shiny, gelatinous, minutely radially rugulose, ± pruinose in places, at first dull blue at apex, below apex, becoming dull blue towards margin; margin decurved, entire, sulcate, striate, faintly translucent-striate. |
|
White, thin, fragile. | White, fragile, thin. | Thin, pallid, pliant. | – | – | Very thin to moderately thick at apex, translucent white or translucent greyish white. |
|
16–28 reaching the stem, adnate to slightly adnexed with a short tooth, narrowly spaced, white, with intervenose veins. | 14–25 reaching the stem, adnate to slightly adnexed with a short tooth, white, with unconspicuous intervenose veins, edges concolorous with the face. | Close to crowded, 18–25 reach the stipe, two or three tiers of lamellulae, ascending-adnate, sometimes narrowly adnate or practically free, narrow to moderately broad, white or tinged grayish, edges slightly fimbriate. | 17–25 reaching the stem, ascending, adnexed, greyish to greyish brown; edge whitish, at times yellowish, greenish or bluish near the cap margin. | 9–14 reaching the stem, ascending, adnexed to fairly broadly adnate or almost free, sometimes with a pseudocollarium, whitish or pale grey; edge whitish and separable as an elastic-tough thread. | Free from stipe or adnately attached to obvious circular descent of pileal flesh, moderately close to distant, five to seven per quadrant, subventricose, moderately broad to broad; edge marginate, blue; sides minutely pruinose, white; with one or two series of lamellulae. |
|
48–76 × 1.5–2.0 mm, equal or slightly broadened below, hollow, fragile, pruinose, white, base acid blue in the whole age, covered with white fibrils. | 32–46 × 1.0–2.0 mm, equal, base sometimes slightly broaden, fragile, hollow, pruinose, puberulous entirely when young, becoming sparely especially in the middle part when old, yellowish brown to light brown, base with acid blue tinge, covered with a bit white fibrils. | 3–8 cm long, 1–2 (2.5) mm. thick, equal, terete, flexuous or strict, tubular, cartilaginous, elastic, at first densely pruinose or minutely pubescent over all form a dense coating of caulocystidia, somewhat glabrescent, base mycelioid, the mycelium blue at first but soon fading to white, bluish to greenish blue above at first, soon fading to grayish or finally sordid brownish. | 40–70 × 0.5–2 mm, cylindrical, entirely covered with a dense and fairly coarse, white pubescence, greyish brown, usually somewhat paler at the apex, occasionally with a slight lilaceous or violaceous tint; base at times somewhat rooting, concolorous or with some blue-green stains or entirely blue, even the substrate may be stained blue. | 5–30 (–70) × 0.5–1 mm, cylindrical, entirely puberulous, glabrescent in the middle part, pale grey to hyaline-white; base hirsute, sky blue (also in the flesh), springing from a patch of fine, radiating, white fibrils. | Up to 22 mm long, cylindrical, moist to dry, often pruinose especially towards base, translucent white, attached to substratum via white pruinose disc borne on a flattened dull blue base. |
|
Indistinctive | Indistinctive | Mild | Indistinct to raphanoid. | Smell none or reported as faintly nitrous; taste not recorded. | Odour not distinctive. |
|
(9.0) 9.3–11.6 (11.8) × (6.0) 6.2–7.3 (7.7) μm, Q = 1.5–1.7, ellipsoid, amyloid. | (6.2) 6.4–7.7 (7.9) × (4.4) 4.7–5.8 (6.0) μm, Q = 1.23–1.54, broadly ellipsoid to ellipsoid, amyloid. | 6–8 × 6–7 (8) μm, globose or subglobose, amyloid. | 7.5–10.7 × 4.5–6 μm, Q = 1.5–1.9, Qav ≈ 1.6, pip-shaped, amyloid. | 6.5–9 × 4–5 μm, Q = 1.6–2.2, Qav ≈ 1.8, pip-shaped, amyloid. | (54/3), 8.4–11.6 (x̄ = 9.9, SD = ± 0.7) × 5.7–8.8 (x̄ = 7.0, SD = ± 0.6) μm, Q = 1.4, broadly ellipsoidal rarely subglobose, with prominent short, oblique apiculus, amyloid. |
|
22–29 × 7–9 μm, 4- or 2-spored. | 26–35 × 6–12 μm, 4- or 2-spored. | 4-spored | 30–40 × 6–7 μm, clavate, 4-spored. | 18–25 × 6.5–11 μm, clavate, 4-spored. | (27/2), 21.6–39.8 (x̄ = 29.0, SD = ± 5.2) × 8.3–16.0 (x̄ = 11.6, SD = ± 2.6) μm, 4-spored, rarely 2-spored, sterigmata to 8.8 μm long. |
|
40–62 × 4–6 μm, clustered, abundant, long clavate or cylindrical, apically broadly rounded, thin-walled, hyaline, forming a sterile lamellae edge. | 32–48 × 4–6 μm, abundant, clustered, cylindrical or long clavate, apically broadly rounded, thin-walled, hyaline, forming a sterile lamellae edge. | Abundant, 32–60 × 5–8 μm, subfusoid with obtuse apices but becoming more or less cylindric, sometimes flexuous, smooth, hyaline. | 16–45 × 3.5–7 μm, clavate, subfusiform or more often cylindrical. | 9–20 × 5.5–7 μm, embedded in gelatinous matter, clavate to obpyriform, with few, simple to branched excrescences, 3–14 × 1–1.5 μm. | Abundant, (30/1), 16.8–44.8 (x̄ = 25.5, SD = ± 6.55) × 5.6–13.6 (x̄ = 8.4, SD = ± 1.8) μm, filamentous, cylindrical, clavate to ovoid, sometimes ventricose at base, with nodulose excrescences. |
|
Absent | Absent | Not differentiated | Absent | Absent | Absent |
|
Hyphae 1–4 μm wide, sparse, smooth or sparsely coated with simple, cylindrical excrescences or inflated cells, 3.1–11.2 × 0.8–1.7 μm, embedded in gelatinous matter. | Hyphae 2–4 μm wide, with simple, cylindrical excrescences, 2.0–6.4 × 0.6–1.8 μm, embedded in gelatinous matter. | A thick gelatinous pellicle (blue color located along the surface of the pellicle in incompletely gelatinized hyphae). | Hyphae 2–4.5 μm wide, branched, anastomosing, smooth with scattered, cylindrical excrescences, embedded in a layer of gelatinous matter. | Hyphae 1.5–3.5 μm wide, embedded in gelatinous matter, very branched, covered with scattered, simple to branched excrescences, protruding through the gelatinous layer. | Hyphae (28/1), 2.8–8.0 (x̄ = 5.4, SD = ± 1.4) μm in diam., nodulose-diverticulate with dense nodulose to cylindrical excrescences, gelatinized. |
|
Hyphae 3–8 μm in diameter, smooth, hyaline. | Hyphae 3–6 μm in diameter, smooth, hyaline. | – | Hyphae 2–3.5 μm wide, smooth | Hyphae 1–3 μm wide, smooth. | Hyphae (26/1), 1.6–3.2 (x̄ = 2.4, SD = ± 0.4) μm in diam., not gelatinized. |
|
38–69 × 6–8 μm, long cylindrical, smooth, transparent. | 19–40 × 4–8 μm, smooth, transparent, two shapes: fusiform or cylindrical. | Covered with numerous cystidia, elongated and flexuous. | 50–145 × 8–11.5 μm, fusiform to subcylindrical. | Up to 60 × 7 μm, simple to furcate or somewhat branched. | Often fasciculate, (25/1), 50.6–128.0 (x̄ = 75.0, SD = ± 19.8) × 5.0–8.8 (x̄ = 6.3, SD = ± 1.1) μm, filamentous to slightly ventricose especially towards base, rarely bifurcate. |
|
Present | Present | Present | Present | Present | Present |
|
Scattered, on humus and fallen leaves in |
Scattered, on rotten wood in |
Single, scattered or gregarious on debris, decaying wood, or on the bark around the bases of live trees of oak in particular, but also occurring quite frequently on decaying wood of basswood, elm, beech, and other hardwoods. | On wood and woody debris, mostly from conifers but also deciduous trees, also among leaves and needles. | On conifers ( |
Generally gregarious, often abundant, rarely solitary or scattered, on fallen decayed logs or stumps of |
|
China | China | North America (Alabama, Carolina, New York, Tennessee, Pennsylvania, Michigan); Canada (Nova Scotia, Ontario, Manitoba) | Europe (Scandinavia, Netherlands, Italy) | Europe (UK, Denmark, Italy) | Australia and New Zealand |
|
Summer to autumn. | Late summer to early autumn. | Spring to fall, more abundant locally in the spring. | Late summer to late autumn, rarely in spring. | Summer to autumn. | March to July. |
|
This study | This study |
|
Microscopic features of
Morphological features of
This species is characterised by dark brown pileus with a blue margin and the stipe densely pruinose, entirely covered with puberulous hairs and stipe basal disc and acanathocysts of pileipellis absent.
China. Jilin Province: Chixi Protection Station, Erdaobaihe Town, Antu County, Yanbian Korean Autonomous Prefecture,
Refers to the pileus, which is blue at the margin.
Pileus 3.5–13 mm in diameter, parabolic, obtusely conical when young, hemispherical, campanulate with age, with an umbo at the centre, shallowly sulcate, translucently striate, smooth, slightly gelatinous, the margin infrequently out of flatness, dark brown (6F5–6F7), disc brown (6E6–6E7), becoming greyish-blue (23B5) to blue (23B7) towards the margin (Fig.
Fresh basidiomata of
Basidiospores [60/3/2] (6.2) 6.4–
Morphological features of
Scattered on rotten wood in
Microscopic features of
Jilin Province, China.
Jilin Province: Hancongling, Erdaobaihe Town, Antu County, Yanbian Korean Autonomous Prefecture,
The diagnostic features of
1 | Cheilocystidia non-smooth |
|
– | Cheilocystidia smooth (sect. |
|
2 | Acanthocysts present (sect. |
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– | Acanthocysts absent |
|
3 | Stipe with basal disc (sect. |
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– | Stipe without basal disc (sect. |
|
4 | Basidiospores subglobose |
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– | Basidiospores broadly ellipsoid to ellipsoid |
|
5 | Caulocystidia of two types: (1) fusiform or cylindrical, 19–40 × 4–8 μm; (2) extremely long, cylindrical (length > 100 μm) |
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– | Caulocystidia of one type, fusiform, subcylindrical to cylindrical (length < 100 μm) |
|
6 | Pileus pale grey-brown or pale sepia brown, sometimes with an olivaceous, greenish or bluish-green shade; margin often bluish-green or rarely dingy citrine to ochraceous yellow |
|
– | Pileus sky blue, greyish-blue with age; margin blue when young, turning bluish-grey when old |
|
With their blue pileus and gelatinous pileipellis, the new taxa
Although pileus colour has been used as a basis for sectional division in
This study was supported by the Natural Science Foundation of Shandong Province (grant No. ZR2020QC001), the National Natural Science Foundation of China (grant No. 3190012), the Natural Science Foundation of Shandong Province (grant No. ZR2019PC028) and the Innovation Team of Shandong Agricultural Industry Technology System (grant No. 26, SDAIT-07-03). We are extremely grateful for the assistance of the Herbaria of Cornell University and the University of Tennessee and especially appreciate the kind help of Collections Manager Margaret Oliver, Dr P. Brandon Matheny and Curator Teresa Iturriaga with specimen loan requests. We sincerely thank Dr Jianwei Liu for help with phylogenetic analyses and Dr Xiaojuan Deng, Mr Bai Wang, Mr Chi Yang and colleagues of the Guyuan Branch Institute of Ningxia Academy of Agriculture and Forestry Sciences for help with fieldwork. We also thank the editors and reviewers for their corrections and suggestions to improve our work.