Figures (67)  Tables (4)

    • Figure 1. 

      Bayesian Inference (BI) phylogeny of Leucoagaricus and Leucocoprinus s.l. inferred from the concatenated ITS-nrLSU-rpb2-tef1-α dataset. Branch lengths are not drawn to scale for visual clarity. Selected macro- and micro-morphological diagnostic characters are mapped alongside the topology.

    • Figure 2. 

      Partial view of the Bayesian Inference (BI) phylogeny of Leucoagaricus and Leucocoprinus s.l. inferred from the concatenated ITS-nrLSU-rpb2-tef1-α dataset, focusing on Leucoagaricus to illustrate its internal phylogenetic structure and morphological congruence. Due to its size, the comprehensive phylogenetic tree is provided in Supplementary File 2. Selected macro- and micro-morphological diagnostic characters are mapped alongside the topology.

    • Figure 3. 

      Partial view of the Bayesian Inference (BI) phylogeny of Leucoagaricus and Leucocoprinus s.l. inferred from the concatenated ITS-nrLSU-rpb2- tef1-α dataset, focusing on Leucocoprinus s.l. to illustrate its internal phylogenetic structure and morphological congruence. Due to its size, the comprehensive phylogenetic tree is provided in Supplementary File 2. Selected macro- and micro-morphological diagnostic characters are mapped alongside the topology. For the pileus centre covering columns, data in the 'General pileus covering' column are based on original protologues and reliable literature; under 'Surface covering at pileus', '++' indicates explicitly observed microstructures, while '+' indicates character states cautiously inferred from original descriptions.

    • Figure 4. 

      Representative microstructures of Type I at the pileus center. Type I-A cutis (a), (e) Leucoagaricus rutiloides (FJAU77871); (c) Leucocoprinus acystidiatus (FJAU78090). Type I-B interwoven plagiotrichoderm (b) Leucocoprinus fuscannulatus (FJAU78013); (d), (f) Leucocoprinus parviceps (FJAU78046). Scale bars = 50 μm.

    • Figure 5. 

      Representative microstructures of Type II at the pileus center. (a), (b) Type II-A trichoderm. Leucocoprinus bicolorannulatus (FJAU78055); (c), (d) Type II-B catenulate trichoderm. Leucocoprinus guttifer (FJAU78018). Scale bars = 50 μm.

    • Figure 6. 

      Representative microstructures of Type III at the pileus center. Type III-A conioderm. (a), (b) Leucocoprinus parvicanus (FJAU78110); (c), (d) Type III-B moniliform plagiotrichoderm. Leucocoprinus umbrosquamosus (FJAU78072). Scale bars = 50 μm.

    • Figure 7. 

      Basidiomata of Leucoagaricus rubrotinctus complex. La. lateritiopurpureus (a) FJAU77772, (b) FJAU77771. La. rutiloides (c) FJAU77870, (d) FJAU77871, (e) FJAU77869. La. rubrovelutinus (f) FJAU77847, (g) FJAU77846. La. cuscutoides (h) FJAU77946, (i), (j) FJAU77945. La. rutilofbrillosus (k) FJAU77801, (l) FJAU77802. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 8. 

      Basidiomata of Leucoagaricus rubrotinctus complex. La. proximus (a) FJAU77767, (b) FJAU77765. La. rubrobrunneus (c) FJAU77780, (d) FJAU77784, (e) FJAU77783. (f), (g) La. vassiljevae (FJAU77843). (h) La. aurantioruber (FJAU77776). (i) La. subpurpureolilacinus (FJAU77943). (j) La. subnivalis (FJAU77858). (k) La. testaceumbonatus (FJAU77913). (l) La. bulbosus (FJAU77942). Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 9. 

      Leucoagaricus cuscutoides. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 10. 

      Leucoagaricus rutilofibrillosus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 11. 

      Leucoagaricus rutiloides. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 12. 

      Leucoagaricus proximus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e )= 50 μm.

    • Figure 13. 

      Leucoagaricus rubrobrunneus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 14. 

      Basidiomata of Leucoagaricus serenus complex. La. albissimus (a) FJAU77928, (b) FJAU78127. La. brunneimutabilis (c) FJAU77891, (d) FJAU77894. La. albofulvescens (e) FJAU77861, (f) FJAU77863, (g) FJAU77897. La. albofibrillosus (h) FJAU78277, (i) FJAU78276. La. leucotrichus (j), (k) FJAU77805, (l) FJAU77807. Scale bars = l cm. These pictures are copyright of the collectors of specimens.

    • Figure 15. 

      Basidiomata of Leucoagaricus serenus complex. La. subleucotrichus (a), (b) FJAU78135, (c) FJAU77808. La. microsporus (d) FJAU77889, (e) FJAU77887, (f) FJAU77885. La. submicrosporus (g) FJAU77919, (h) FJAU77920. La. centrobrunneolus (i) FJAU77909, (j) FJAU77908. La. albosquamosus (k) FJAU78148, (l) FJAU78140. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 16. 

      Basidiomata of the Leucoagaricus serenus complex. La. magnialbus (a) FJAU77822, (b) FJAU77820. La. similis (c) FJAU77905, (d), (e) FJAU78099. (f), (g) La. aff. similis (FJAU77903). La. pallidoumbonatus (h) FJAU77878, (i) FJAU77876, (j) FJAU77875. La. jianensis (k) FJAU77937, (l) FJAU77936. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 17. 

      Basidiomata of the Leucoagaricus serenus complex. La. ruiliniae (a) FJAU77884, (b), (c) FJAU77880. La. aff. serenus 1 (d) FJAU77789, (e) FJAU77790, (f) FJAU77787. La. aff. serenus 2 (g) FJAU77794, (h) FJAU77792. La. candidus (i) FJAU77754, (j) FJAU77753. (k), (l) La. subcandidus (FJAU77940). Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 18. 

      Leucoagaricus albofibrillosus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 19. 

      Leucoagaricus jianensis. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 20. 

      Leucoagaricus leucotrichus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c)–(e) = 20 μm.

    • Figure 21. 

      Leucoagaricus magnialbus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 22. 

      Leucoagaricus microsporus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 23. 

      Leucoagaricus similis. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 24. 

      Leucoagaricus subleucotrichus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 25. 

      Leucoagaricus aff. serenus 1. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 26. 

      Leucoagaricus aff. serenus 2. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 27. 

      Leucoagaricus submicrosporus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 28. 

      Leucoagaricus aff. similis. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 29. 

      Basidiomata of Leucoagaricus species. La. luteus (a) FJAU77804, (b) FJAU77803. La. centriluteus (c), (d) FJAU77798, (e) FJAU77797. La. conifericola (f) FJAU77933, (g) FJAU77932, (h) FJAU77935). La. subcrystallifer (i) FJAU77834, (j) FJAU77831). La. atroviridis (k) FJAU77840, (l) FJAU77842. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 30. 

      Leucoagaricus centriluteus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 31. 

      Leucoagaricus conifericola. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 32. 

      Leucoagaricus luteus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 33. 

      Basidiomata of Leucocoprinus species. Lc. acystidiatus (a) FJAU78091, (b), (c) FJAU78090. (d), (e) Lc. crassibasis (FJAU78107). Lc. fuscannulatus (f), (g) FJAU78014, (h) FJAU78010, (i) FJAU78009. Lc. griseocentralis (j) FJAU78118, (k) FJAU78006, (l) FJAU78003. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 34. 

      Basidiomata of Leucocoprinus species. (a), (b) Lc. parvicanus (FJAU78110). (c), (d) Lc. porosporus (FJAU78028). Lc. sinumbellatus (e) FJAU77949, (f) FJAU78109). Lc. sphaerodermus (g), (h) FJAU78049, (i) FJAU78050. Lc. subincarnatus (j) FJAU78134, (k), (l) FJAU78095. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 35. 

      Basidiomata of Leucocoprinus species. (a) Lc. subumbrosquamosus (FJAU78071). (b) Lc. umbrosquamosus (FJAU78072). Lc. guttifer (c), (d) FJAU78021, (e) FJAU78018. Lc. floccipes (f) FJAU78078, (g) FJAU78073, (h) FJAU78075, (i) FJAU78074). Lc. helianthoides (j), (k) FJAU78092, (l) FJAU78093. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 36. 

      Basidiomata of Leucocoprinus and Macropsalliota species. Lc. luteoguttatus (a) FJAU78022, (b) FJAU78023, (c) FJAU78026). Lc. ochraceosquamosus (d) FJAU78017, (e), (f) FJAU78016. Lc. sudans (g), (h) FJAU78274, (i) FJAU78106). (j), (k) Map. aff. subtropica (FJAU72225). (l) Map. subtropica. (FJAU77730). Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 37. 

      Basidiomata of Leucocoprinus species. Lc. cinerascens (a) FJAU77966, (b) FJAU77965. (c) Lc. birnbaumii (FJAU79140). (d) Lc. licmophorus (FJAU79141). Lc. lugdunensis (e) FJAU77961, (f) FJAU77962. (g) Lc. nympharum (i) FJAU77969). (h) Lc. tangerinus (FJAU78137). Lc. parviceps (i) FJAU78047, (j), (k) FJAU78046. (l) Lc. lacteus (FJAU78083). Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 38. 

      Leucocoprinus acystidiatus. (a) Basidiomata. (b) Basidia. (c) Basidiospores. (d) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 39. 

      Leucocoprinus crassibasis. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 40. 

      Leucocoprinus fuscannulatus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 41. 

      Leucocoprinus griseocentralis. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 42. 

      Leucocoprinus parvicanus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 43. 

      Leucocoprinus porosporus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 44. 

      Leucocoprinus sinumbellatus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 45. 

      Leucocoprinus sphaerodermus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 46. 

      Leucocoprinus subincarnatus. (a) Basidiomata. (b) Basidia. (c) Basidiospores. (d) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 47. 

      Leucocoprinus subumbrosquamosus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 48. 

      Leucocoprinus umbrosquamosus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 49. 

      Leucocoprinus floccipes. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d), (f) = 20 μm; (e) = 50 μm.

    • Figure 50. 

      Leucocoprinus guttifer. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d), (f) = 20 μm; (e) = 50 μm.

    • Figure 51. 

      Leucocoprinus helianthoides. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d), (f) = 20 μm; (e) = 50 μm.

    • Figure 52. 

      Leucocoprinus luteoguttatus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d), (f) = 20 μm; (e) = 50 μm.

    • Figure 53. 

      Leucocoprinus ochraceosquamosus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e), (f) = 50 μm.

    • Figure 54. 

      Leucocoprinus sudans. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d), (f) = 20 μm; (e) = 50 μm.

    • Figure 55. 

      Leucocoprinus lacteus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 56. 

      Leucocoprinus parviceps. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 57. 

      Basidiomata of Leucocoprinus species. Lc. comatus (a) FJAU78115, (b) FJAU78065. Lc. rutilobrunneus (c) FJAU78029, (d) FJAU78032, (e) FJAU78030, (f) FJAU78036. Lc. bicolorannulatus (g) FJAU78054, (h), (i) FJAU78055, (j) FJAU78053, (k), (l) FJAU78132. Scale bars = 1 cm. These pictures are copyright of the collectors of specimens.

    • Figure 58. 

      Basidiomata of Leucocoprinus species. Lc. persicinus (a) FJAU78037, (b) FIAU78039. Lc. caeruleipes (c), (d) FJAU78045, (e) FJAU78044. Lc. versicolor (f) FJAU78041, (g), (h) FJAU78042. Lc. caffeaceus (i) FJAU78103, (j) FJAU78104. Lc. transeptatus (k) FJAU78067, (l) FJAU78069. Scale bars = l cm. These pictures are copyright of the collectors of specimens.

    • Figure 59. 

      Dried specimens of Leucocoprinus and Macropsalliota species. (a) Lc. comatus (left two: FJAU78115, right two: FJAU78065). (b) Lc. rutilobrunneus (FJAU78029). (c) Lc. bicolorannulatus (from left to right: FJAU78051, FJAU78052, FJAU78053, FJAU78054, and FJAU78055). (d) Lc. persicinus (FJAU78037). (e) Lc. caeruleipes (left two basidiomata photographed on 24 May 2025, right two photographed on 6 September 2025; first and third: FJAU78044, second and fourth: FJAU78045). (f) Lc. versicolor (left two basidiomata photographed on 24 May 2025, right two photographed on 6 September 2025; left three: FJAU78043, right one: FJAU78041). (g) Lc. caffeaceus (FJAU78103). (h) Lc. transeptatus (left one: FJAU78068, right two: FJAU78067). (i) Map. aff. subtropica (FJAU72226). These pictures are copyright of X.Y. Zhou.

    • Figure 60. 

      Leucocoprinus bicolorannulatus. (a) Basidiomata. (b) Surface covering at pileus centre. (c) Basidiospores. (d) Cheilocystidia. (e) Basidia. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 61. 

      Leucocoprinus comatus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 62. 

      Leucocoprinus persicinus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Pileus covering at pileus centre. (f) Stipe covering. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (f) = 20 μm.

    • Figure 63. 

      Leucocoprinus rutilobrunneus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 64. 

      Leucocoprinus caeruleipes. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 65. 

      Leucocoprinus caffeaceus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 66. 

      Leucocoprinus transeptatus. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Figure 67. 

      Leucocoprinus versicolor. (a) Basidiomata. (b) Basidiospores. (c) Basidia. (d) Cheilocystidia. (e) Surface covering at pileus centre. Scale bars: (a) = 1 cm; (b) = 10 μm; (c), (d) = 20 μm; (e) = 50 μm.

    • Author (year) Methodological basis Key contribution and diagnostic features
      Patouillard (1888)[5] Morphology Established Leucocoprinus to segregate white-spored species from Lepiota (Pers.) Gray that exhibit Coprinus-like characteristics and possess a distinct germ pore.
      Locquin (1945)[6] Morphology Subdivided Leucocoprinus into four subgenera: subgen. Eu-Leucocoprinus (Pat.) emend., subgen. Leucoagaricus Locq., subgen. Hiatula (Fr.) emend., and subgen. Leucobolbitius (Lange) emend.
      Singer (1948a)[8] Macro- and micromorphology Elevated Leucocoprinus subgen. Leucoagaricus (Locq.) to the generic rank: Leucoagaricus. Characterized by small, colorless, dextrinoid, and metachromatic spores with a germ pore, and the presence of an annulus (Type: La. macrorhizus [Locquin] Singer= Lc. barssii [Zeller] Migl. & Donato). Divided into two sections: – sect. Macrospori: spores 12–15 µm (Type: La. excoriatus = Macrolepiota excoriata [Schaeff.] Wasser). – sect. Rubrotincti: spores < 12 µm, with brightly colored pileus squamules (Type: La. rubrotinctus [Peck] Singer).
      Singer et al. (1972)[9] Macro- and micromorphology Further expanded the infrageneric classification of Leucoagaricus by establishing four sections: – sect. Sphaerocystophori: pileipellis composed of sphaerocysts (Type: La. exannulatus Singer). – sect. Sculpturati: similar to sect. Rubrotincti, but with ornamented spores (Type: La. rubrosquamosus [Rick] Singer). – sect. Piloselli (Kühn.) Singer: Lamellae becoming pinkish at maturity; context often turning red/darkening; spores smooth, lacking a germ pore (Type: La. georginae [W.G. Sm.] Singer= Lc. georginae [W.G. Sm.] M. Asif, Saba & Vellinga). – Sect. Annulati (Fr.) Singer: Species with smooth, dextrinoid spores possessing a germ pore; often robust basidiomata (Type: La. naucinus [Fr.] Singer = Lc. leucothites [Vittad.] Redhead).
      Singer (1986)[10] Macro- and micromorphology The tribe Leucocoprineae Sing. was defined to contain seven genera: Clarkeinda, Chlorophyllum, Macrolepiota, Volvolepiota, Sericeomyces, Leucoagaricus, and Leucocoprinus.
      Re-evaluated Leucoagaricus and Leucocoprinus. Expanded Leucoagaricus to six sections by adding sect. Leucoagaricus (Type: La. macrorhizus Locq. ex Singer = Lc. barssii [Zeller] Migl. & Donato) and sect. Rubrotincti Singer, which includes species with non-brightly colored, fibrous, scaly, or tomentose pileus coverings (often colorless or sparsely pigmented), and smooth spores.
      Candusso & Lanzoni (1990)[15] Macro- and micromorphology Reviewed the infrageneric classification. Removed subgen. Hiatula and subgen. Leucobolbitius from Leucocoprinus, restricting it to two sections: – sect. Leucocoprinus: slender basidiomata, thin pileus scales, small, elliptical basidiospores (Type: Lc. cepistipes [Sowerby] Pat.). – sect. Denudali Herink: pileus scales with sphaerocysts; spores with inconspicuous or immature germ pores (Type: Lc. denudatus [Sacc.] Singer = Lc. straminellus [Bagl.] Narducci & Caroti).
      Maintained six sections in Leucoagaricus, largely consistent with Singer[10], but replaced sect. Sphaerocystophori with sect. Pulverulenti Bon.
      Akers (1997)[16] Macro- and micromorphology Recognizing four sections within Leucocoprinus: – sect. Denudali Herink. – Sect. Velutipedes Heinem.: basidiospores with a distinct apical pore; stipe base with scales; pileipellis lacking sphaerocysts and non-hymeniform. – Sect. Leucocoprinus (Pat.) Heinem. – Sect. Pepinispori Heinem.: basidiospores flattened-amygdaliform; distinct apical pore absent or incomplete.
      Maintained the sectional views of Candusso & Lanzoni[15] for Leucoagaricus.
      Migliozzi & Testoni (2000)[35] Macro- and micromorphology Established the new sect. Cystidiosi Migliozzi et Testoni (Type: La. pleurocystidius). Characterized by pleurocystidia (present in the subhymenium, but lacking clamp connections).
      Vellinga (2004)[17] ITS and nrLSU phylogeny Demonstrated that Leucoagaricus, Leucocoprinus, and Sericeomyces together form a single, large monophyletic clade, revealing that these genera are not monophyletic individually under traditional morphological concepts.
      Vellinga et al. (2011)[18] Multi-locus phylogeny (ITS, nrLSU, rpb2, tef1) Revealed that tef1 sequences resolve the Leucoagaricus/Leucocoprinus complex into distinct clades corresponding to traditional sections (e.g., La. sect. Rubrotincti, La. sect. Piloselli). Furthermore, demonstrated that the La. americanus clade is phylogenetically distant from the core complex and allied with Micropsalliota
      Redhead (2023)[22] Nomenclatural reassessment Proposed several new combinations by transferring species from related genera into Leucocoprinus to address taxonomic issues.
      Asif et al. (2024)[23] ITS and nrLSU phylogeny Proposed 64 new combinations within Leucoagaricus and Leucocoprinus.
      Migliozzi et al. (2024)[24] Nomenclatural reassessment Transferred sect. Pilosellae and sect. Rubrotincti from Leucoagaricus to Leucocoprinus; proposed 34 new combinations.
      Yang et al. (2024)[25] Multi-locus phylogeny (ITS, nrLSU, rpb2, tef1) Erected Candelolepiota and Macropsalliota; redefined Leucoagaricus s.s. to include only original sect. Leucoagaricus and Sericeomyces.
      Li et al. (2025)[34] Multi-locus phylogeny and comprehensive morphological examination Expanded species-level diversity by describing 10 new taxa. Highlighted that the circumscription of the revised Leucocoprinus s.l. remained unsettled and identified a distinct 'Clade Cretaceous' separate from the core Leucocoprinus.
      Radnóti et al. (2025)[26] Multi-locus phylogeny (ITS, nrLSU, rpb2, tef1) Erected Mystagaricus to accommodate La. brunneolilacinus, positioning it outside the core Leucoagaricus and Leucocoprinus s.l. group.
      Li et al. (2025)[29] Phylogenomics (ITS, nrLSU, rpb2, tef1) Erected the new genus Conioexocarpus and the new subfamily Leucocoprinoideae to accommodate Leucoagaricus, Leucocoprinus, Conioexocarpus, Candelolepiota, Macropsalliota, and Micropsalliota.
      Yang et al. (2026)[30] Multi-locus phylogenetic analyses (ITS, nrLSU, rpb2, tef1-α), coalescent-based species tree estimation (ASTRAL), and comprehensive microstructural investigations Restructured the generic framework of subfam. Leucocoprinoideae. Established the new genera Pulchrolepiota and Tristolepiota to resolve the paraphyly within Leucocoprinus s.l. Defined Leucocoprinoideae as comprising five genera (Conioexocarpus, Leucoagaricus, Leucocoprinus s.s., Pulchrolepiota, and Tristolepiota) based on pileus center microstructures (trichoderm, conioderm, or cutis) and the presence/absence of hymenial physalides. Additionally established subfam. Micropsalliotoideae to accommodate Mesopsalliota (gen. nov.), Macropsalliota, and Candelolepiota.
      Vellinga et al. (2026)[31] Nomenclatural reassessment Facilitated a broadly defined Leucocoprinus concept by proposing 10 new combinations for North American taxa. Emphasized that generic delimitation should integrate practical taxonomic utility and broader criteria, rather than relying exclusively on phylogenetic topological parameters.

      Table 1. 

      Chronological summary of key taxonomic revisions and framework shifts in Leucoagaricus and Leucocoprinus s.l.

    • TaxonBasidiaCheilocystidia
      Lc. sabinae[88]Basidia tetra-sterigmate, clavate, or cylindrical with a square base, 18–30 × 6.5–10 µm.Cheilocystidia 13.5–21 × 3–6 µm, cylindrical-capitulated, some with flexuous outline, infrequent and unevenly distributed along the lamella edge.
      Lc. subhymenoderma[87]Basidia 20–30 × 7–9 μm, (1–2–) 4-spored.Cheilocystidia 25–35 × 7–12 μm, clavate to ± ventricose, or sometimes constricted.
      Lc. pegleri[61]Basidia 18–23 × 6–8 μm, clavate, tetrasporic.Cheilocystidia 12–23 × 7–11 μm, clavate, basidia-like, not very visible on the lamellar edge
      Lc. roseovertens[61]Basidia 14.5–24.5 × 6–9 μm, clavate, tetrasporic.Cheilocystidia 12–23 × 7–13 μm, ventricose, clavate, hyaline, emerging in little tufts from the lamellar edge.
      Lc. acystidiatus (this study)Basidia 13–24 × 5–8 μm, clavate, 4(–2)-spored.No specialized terminal hyphal cells observed.
      Lc. subincarnatus (this study)Basidia 16–22 × 6–9 μm, clavate, 4(–2)-spored.No specialized terminal hyphal cells observed.

      Table 2. 

      Leucocoprinus species possessing poorly developed cheilocystidia.

    • 1 Habitat restricted to sandy soils; basidiomata almost entirely buried in sand Lc. taniae
      1' Habitat not sandy soils; basidiomata epigeous 2
      2 Habitat strictly linked to attine ants Lc. karjaticus
      2' Habitat not linked to attine ants 3
      3 Growing on decaying wood Lc. dacrytus
      3' Growing on humus or soil 4
      4 Basidiospores possessing a distinct germ pore 5
      4' Basidiospores lacking a germ pore 7
      5 Pseudoparaphyses absent Candelolepiota sinica
      5 Pseudoparaphyses present 6
      6 Cheilocystidia 35–48 × 12–22 μm, clavate to broadly fusiform, with papillate or capitate apices Lc. lugdunensis
      6 Cheilocystidia 25–60 × 8–15 μm, clavate, with lageniform apices Lc. cepistipes
      7 Droplets secreted from the entire basidioma when fresh 8
      7 Droplets secreted exclusively from the stipe when fresh 15
      8 Currently known from Europe or the Americas 9
      8 Currently known only from Asia (China) 11
      9 Distributed in Europe; lamellae slightly greyish-white, sometimes with a pinkish sheen; cheilocystidia frequently with peculiar apical appendages Lc. tener
      9 Distributed in the Americas; lamellae white to yellowish; cheilocystidia without peculiar apical appendages 10
      10 Cheilocystidia 22–58 × 7–20 μm, clavate, lacking mucronate, lageniform, or bifurcated apices Lc. stillatus
      10 Cheilocystidia 24–74 × 8–30 μm, clavate, mostly mucronate, lageniform, or bifurcated Lc. margaritifer
      11 Stipe surface lacking conspicuous squamules 12
      11 Stipe surface covered with squamules 13
      12 Basidiospores 6.1–7.1 × 3.6–4.2 μm (avl × avw = 6.4 × 3.8 μm); cheilocystidia narrowly utriform to clavate Lc. luteoguttatus
      12 Basidiospores generally smaller, 5.2–6.6 × 2.9–3.4 μm (avl × avw = 5.8 × 3.2 μm); cheilocystidia fusiform to oblong Lc. sudans
      13 Cheilocystidia subfusiform to clavate, with distinct apical appendages Lc. shixingensis
      13 Cheilocystidia narrowly cylindrical, narrowly fusiform, or clavate, lacking apical appendages 14
      14 Stipe surface below the annulus covered from the mid-region to the base with golden-brown to gingerbread-brown floccose squamules Lc. floccipes
      14 Stipe surface below the annulus off-white, yellowish-white, or brownish-yellow with fine concolorous squamules Lc. flavirobustus
      15 Pileus surface glabrous to finely fibrillose, lacking distinct squamules Lc. glaber
      15 Pileus surface with distinct coloured squamules 16
      16 Pileus squamules pale gold, golden-brown, or greyish-brown 17
      16 Pileus squamules pale orange, brownish-yellow, reddish-brown, or orange-red 18
      17 Mature pileus relatively large, 2.7–3.5 cm in diam Lc. guttifer
      17 Mature pileus relatively small, 1.1–1.5 cm in diam Lc. ochraceosquamosus
      18 Currently known only from India Lc. brunneodiscus
      18 Currently known only from China 19
      19 Basidiospores 5.2–6.3 × 3.2–4.1 μm Lc. helianthoides
      19 Basidiospores 6.5–7.0 × 4.0–4.5 μm Lc. tangerinus
    • 1 Basidiomata turning blue, green, blue-green, or brown upon bruising 2
      1' Basidiomata not discolouring upon bruising 9
      2 2 Basidiomata turning brown or darkening upon bruising 3
      2' Basidiomata turning blue or green upon bruising 4
      3 Stipe yellow, turning brown upon bruising, and transitioning to dark blue upon desiccation; cheilocystidia 17.2–28.6 × 8.7–11.7 μm Lc. petasatus
      3' Basidiomata darkening more or less upon bruising, transitioning to blue-green upon desiccation; cheilocystidia 26–43 × 9.5–22 μm, variably clavate Lc. vernalis
      4 Basidiomata turning blue upon bruising 5
      4' Basidiomata turning green upon bruising 7
      5 Cheilocystidia relatively small, 11.5–17 × 9–10.6 μm, pyriform to sublageniform Lc. phaeocyaneus
      5' Cheilocystidia relatively large, 10–64.3 × 7.5–21 μm, clavate, narrowly clavate, broadly clavate, pyriform, or sometimes sphaeropedunculate 6
      6 Cheilocystidia 40.5–64.3 × 10.9–16.8 μm, clavate, narrowly clavate, or broadly clavate Lc. atrocaeruleus
      6' Cheilocystidia 10–50 × 7.5–21 μm, broadly clavate to pyriform, sometimes sphaeropedunculate, rarely clavate Lc. viriditinctus
      7 Basidia characterised by a digitate apical appendage Lc. digitatocystis
      7' Basidia lacking a digitate apical appendage 8
      8 Cheilocystidia narrowly clavate to clavate, sometimes fusiform Lc. virens
      8' Cheilocystidia subcylindrical, broadly clavate to pyriform, rarely clavate Lc. beijingensis
      9 Cheilocystidia absent Lc. beninensis
      9' Cheilocystidia present 10
      10 Cheilocystidia frequently 1- to 2-septate, with the terminal segment distinctly digitate Lc. transeptatus
      10' Cheilocystidia llacking a distinctly digitate terminal segment 11
      11 Basidiomata turning blue then green upon desiccation, but ultimately not acquiring a yellowish hue Lc. actinophorus
      11' Basidiomata turning blue then green upon desiccation, ultimately acquiring a yellowish hue, or exhibiting a highly transient blue phase 12
      12 Pileus surface with distinct brown-red to hickory-brown floccose squamules Lc. caffeaceus
      12' Pileus surface lacking distinct floccose squamules 13
      13 Cheilocystidia 21.5–39 × 10–20 μm, clavate to broadly clavate Lc. coerulescens
      13' Cheilocystidia 27–57 × 5–12 μm, narrowly cylindrical to cylindrical, or narrowly clavate 14
      14 Stipe transiently developing a pale blue hue upon desiccation, before the entire basidioma acquires a greenish tint and ultimately turns yellow Lc. caeruleipes
      14' Stipe completely lacking an initial blue phase upon desiccation; basidiomata directly transitioning to a greenish, and subsequently yellowish, phase Lc. versicolor