Fungus Flora of Tropical Africa

Volume 1

 

 MONOGRAPH OF MARASMIUS, GLOIOCEPHALA, PALAEOCEPHALA AND SETULIPES IN TROPICAL AFRICA

 

by  Vladimír Antonín


Plates / Planches


Contents        

Foreword by J. Rammeloo

Introduction

Material and methods

Characters used for the identification of marasmioid and collybioid taxa

1  Macroscopical characters

  1.1  Basidiocarps

  1.2  Pileus

  1.3  Lamellae

  1.4  Stipe

  1.5  Rhizomorphs and sterile stipes

  1.6  Smell and taste

2  Microscopical characters

  2.1  Structure of the pileipellis

  2.2  Spores

  2.3  Characters of the hymenium

  2.4  Trama

  2.5  Stipitipellis and caulocystidia

  2.6  Gelatinous context

  2.7  Chemical reactions of hyphae

3  Ecological characters

Description of macroscopic characters

Abbreviations

Key to marasmioid and collybioid genera

 

Marasmius Fr.

Brief history of Marasmius collections from tropical Africa

Key to sections of the genus Marasmius

Sect. Marasmius

Key to tropical African species

Species descriptions

Subsect. Marasmius

1. Marasmius louisii Singer

2. Marasmius cupressiformis Berk.

3. Marasmius rotalis Berk. & Broome

  3.1. var. rotalis

  3.2. var. latisporus Antonín

4. Marasmius apatelius Singer

5. Marasmius somalomoensis Antonín

6. Marasmius colorimarginatus Antonín

Subsect. Sicciformes

7. Marasmius subruforotula Singer

8. Marasmius conicopapillatus Henn.

9. Marasmius crinisequi F. Muell.

10. Marasmius curreyi Berk. & Broome var. distantifolius Antonín

11. Marasmius yangambiensis Singer

12. Marasmius aurantiostipitatus Antonín & P. Roberts

13. Marasmius guyanensis Mont.

14. Marasmius lovedalensis Antonín & Verbeken

15. Marasmius nigrobrunneus (Pat.) Sacc.

Sect. Hygrometrici Kühner

Key to tropical African species

Species descriptions

16. Marasmius thwaitesii Berk. & Broome

17. Marasmius minutoides Antonín

  17.1. var. minutoides

  17.2. var. angustisporus Antonín

18. Marasmius nyikae Antonín

19. Marasmius parviconicus Pegler

20. Marasmius mulanjensis Antonín

21. Marasmius subalbidulus Antonín

Sect. Leveilleani Singer

Species descriptions

22. Marasmius leveilleanus (Berk.) Pat.

Sect. Epiphylli Kühner

Synopsis of tropical African species

Subsect. Eufoliatini Singer

Species description

23. Marasmius foliiphilus Antonín

Sect. Fusicystides Singer

Species description

24. Marasmius longicystidiatus Antonín

Sect. Chordales Fr.

Key to tropical African species

Species descriptions

25. Marasmius lolema Beeli

26. Marasmius pegleri Courtec.

27. Marasmius schreursii Antonín

28. Marasmius mvumae Antonín & C. Sharp

Sect. Neosessiles Singer

Key to tropical African species

Species descriptions

29. Marasmius neosessilis Singer

30. Marasmius bururiensis Antonín

31. Marasmius cf. sejunctus Singer

32. Marasmius aff. cecropiae Dennis

33. Marasmius cyphella Dennis & D.A. Reid

Sect. Globulares Kühner

Key to tropical African species

Species descriptions

34. Marasmius arborescens (Henn.) Beeli

35. Marasmius lacteoides Antonín

36. Marasmius albertianus Singer

37. Marasmius kigwenensis Antonín

38. Marasmius favoloides Henn.

39. Marasmius violaceoides Antonín

40. Marasmius mesosporus Singer

41. Marasmius zenkeri Henn.

42. Marasmius bekolacongoli Beeli

43. Marasmius brunneolus (Beeli) Singer

44. Marasmius tshopoensis Antonín

45. Marasmius missangoënsis Pat.

46. Marasmius witteanus Singer

47. Marasmius goossensiae Beeli

48. Marasmius muramwyanensis Antonín

49. Marasmius flavus Singer

50. Marasmius heinemannianus Antonín

51. Marasmius flavidulus Henn.

52. Marasmius latepileatus Antonín & C. Sharp

53. Marasmius albidocremeus Antonín

54. Marasmius staudtii Henn.

  54.1. var. staudtii

  54.2. var. magnisporus Antonín

55. Marasmius camerunensis Antonín & Mossebo

Sect. Sicci Singer

Key to tropical African species

Subsect. Siccini Singer

Series Atrorubentes

Species descriptions

56. Marasmius afrosulphureus Courtec.

57. Marasmius xestocephalus Singer

58. Marasmius xestocephaloides Antonín

59. Marasmius atrorubens (Berk.) Mont.

60. Marasmius subarborescens Singer

61. Marasmius buzungulo Singer

62. Marasmius corrugatiformis Singer

63. Marasmius katangensis Singer

Series Spinulosi

Species descriptions

64. Marasmius mengoënsis Pegler

65. Marasmius setiger Pegler

66. Marasmius jalapensis Murrill

67. Marasmius pseudotorquescens Antonín

68. Marasmius castaneovelutinus Henn.

69. Marasmius fulvovelutinus Beeli

Series Leonini

Species descriptions

70. Marasmius episemus Singer

71. Marasmius ferruginacies Antonín

72. Marasmius leptus Singer

73. Marasmius ochropus Singer

74. Marasmius bubalinus Pegler

75. Marasmius nodulocystis Pegler

76. Marasmius megistus Singer

77. Marasmius lilacinoalbus Beeli

  77.1. var. lilacinoalbus

  77.2. var. lilacinocarmineus Singer

  77.3. var. albus Singer

78. Marasmius striaepileus Antonín

79. Marasmius sierraleonis Beeli

80. Marasmius luteostipitatus Mossebo & Antonín

81. Marasmius carcharus Singer

82. Marasmius haediniformis Singer

83. Marasmius macrolobieti Singer

84. Marasmius conicoparvus Antonín & C. Sharp ad int.

85. Marasmius tanougouensis Antonín

Series Haematocephali

Species descriptions

86. Marasmius haematocephalus (Mont.) Fr.

87. Marasmius longistipitatus Antonín

88. Marasmius robertsii Antonín

89. Marasmius haedinus Berk.

90. Marasmius grandisetulosus Singer

91. Marasmius tenuisetulosus (Singer) Singer

92. Marasmius rubrostipitatus Antonín & P. Roberts

93. Marasmius spegazzinii Sacc. & P. Syd.

94. Marasmius cremeopileatus Antonín & C. Sharp

95. Marasmius pallidopileatus Antonín ad int.

96. Marasmius elaeocephalus Singer

97. Marasmius ferruginoides Antonín

98. Marasmius irangianus Antonín ad int.

99. Marasmius confertus Berk. & Broome

  99.1. var. confertus

  99.2. var. tenuicystidiatus Antonín

  99.3. var. parvisporus Antonín

100. Marasmius strigipes Beeli

101. Marasmius bingaensis Singer

Subsect. Inaequales Singer

Species description

102. Marasmius beelianus Singer

 

Gloiocephala Massee

Key to tropical African species

Species descriptions

1. Gloiocephala albocapitata (Petch) Singer

2. Gloiocephala epiphylla Massee

3. Gloiocephala tezae Antonín

4. Gloiocephala longistipata Antonín ad int.

5. Gloiocephala cf. confusa Singer

6. Gloiocephala lacrimospora Antonín ad int.

7. Gloiocephala mucrocystidiata Antonín

 

Palaeocephala Singer

Species description

8. Palaeocephala cymatelloides (Dennis & D.A. Reid) Singer

 

Setulipes Antonín

Key to tropical African species

Species descriptions

1. Setulipes rhizomorphicola Antonín

2. Setulipes afibulatus Antonín

3. Setulipes brevistipitatus Antonín

4. Setulipes curvistipitatus Antonín

5. Setulipes congolensis (Beeli) Antonín

6. Setulipes kisangensis (Singer) Antonín

7. Setulipes hakgalensis (Petch) Antonín

 

Excluded and doubtful taxa

1. Marasmius albofarinaceus Henn.

2. Marasmius alliacioides Henn.

3. Marasmius allium Eichelbaum

4. Marasmius atroalbus Henn.

5. Marasmius aureus Beeli

6. Marasmius barombiensis Henn.

7. Marasmius baumannii Henn.

8. Marasmius bipindeensis Henn.

9. Marasmius buchwaldii Henn.

10. Marasmius cervinus Henn.

11. Marasmius cinereo-flavidus Henn.

12. Marasmius citrinus Henn.

13. Marasmius crispus Henn.

14. Marasmius cyathula Henn.

15. Marasmius discipes Henn.

16. Marasmius discoideus Henn.

17. Marasmius dulcis Beeli

18. Marasmius dusenii Henn.

19. Marasmius ealaensis Beeli

20. Marasmius elaeicola Henn.

21. Marasmius eligmophyllus De Seynes

22. Marasmius englerianus Henn.

23. Marasmius excentricus Henn.

24. Marasmius fasciculatus Beeli

25. Marasmius ferrugineo-luteus Beeli

26. Marasmius flabellatus Henn.

27. Marasmius friesianus Henn.

28. Marasmius geophyllus Henn.

29. Marasmius gracillimus Henn.

30. Marasmius grandisporus Henn.

31. Marasmius griseoflavus Henn.

32. Marasmius hungo Henn.

33. Marasmius hygrocyboides Henn.

34. Marasmius hymenofallax De Seynes

35. Marasmius jodocodos Henn.

36. Marasmius lilacinostriatus Henn.

37. Marasmius maranguensis Henn.

38. Marasmius matschingili Henn.

39. Marasmius minutulus Henn.

40. Marasmius munsae Henn.

41. Marasmius njalaensis Beeli

42. Marasmius nocticolor De Seynes

43. Marasmius ornatus Henn.

44. Marasmius pahouinensis De Seynes

45. Marasmius pallide-sepiaceus Henn.

46. Marasmius pallidus Henn.

47. Marasmius palmicola Henn.

  var. grisea Henn.

48. Marasmius paradoxus Henn.

49. Marasmius petalocladus De Seynes

50. Marasmius piperatus Beeli

51. Marasmius piperodorus Beeli

52. Marasmius pleurotoides Henn.

53. Marasmius pseudocalopus Henn.

54. Marasmius pseudosplachnoides Henn.

55. Marasmius pygmaeus Henn.

56. Marasmius reniformis Henn.

57. Marasmius reticulatus Henn.

58. Marasmius roseolus Henn.

59. Marasmius rufobrunneus Henn.

60. Marasmius rufus Henn.

61. Marasmius stuhlmannii Henn.

62. Marasmius subcastaneus Henn.

63. Marasmius subcurreyi Henn.

64. Marasmius subimpudicus Henn.

65. Marasmius sublanguidus Henn.

66. Marasmius suboreades Beeli

67. Marasmius subplancus Henn.

68. Marasmius subrhodocephalus Henn.

69. Marasmius subrotula Beeli

70. Marasmius subviolaceus Henn.

71. Marasmius testaceus Henn.

72. Androsaceus thollonis Pat. & Hariot

73. Marasmius togoensis Henn.

74. Marasmius violaceus Henn.

75. Marasmius volkensii Henn.

References

 

Foreword / Avant propos

by J. Rammeloo (Director National Botanic Garden of Belgium) & J. Degreef (Editor)

 

The National botanic garden of Belgium has a long-standing tradition in the study of the mycoflora of central Africa. The oldest collections of Central African Fungi kept in its herbarium are more than a century old (1891, 1895), dating from the period thath Congo was still the private property of the Belgian king Leopold II. The oldest important publication about the mycoflora from Congo is the Reliquiae Dewevreanae (1901) by De Wildeman & Durand.

Since 1923 Mrs M. Goossens-Fontana, a drawing teacher living in the Eala botanic garden (Bas-Congo) started to depict mushrooms using water colour techniques. When returning to Belgium, an amateur mycologist, M. Beeli, stimulated her to collect, to make exsiccatae and spore prints and taught her mushroom microscopy. Afterwards she stayed at Binga and Panzi (Kivu). Her herbarium comprises 1870 numbers and 1331 watercolour drawings, mainly of fleshy fungi, an exceptional tresaure. Since 1926 Beeli regularly published the new findings (about 300 new species), alone or together with Mrs M. Goossens-Fontana in the series Fungi Goossensiani.

In 1935 Beeli convinced the former director of the Brussels botanic garden, W. Robyns, to start the publication of an illustrated mushroom flora of the Congo, the Flore iconographique des champignons du Congo. Seventeen fascicles were published until 1970.

In 1972 P. Heinemann, the new editor, made a new concept for the flora and the title changed to Flore illustrée des champignons d'Afrique centrale. The descriptions became more complete, microscopic characters were added and illustrated. Diagnoses were still published separately, mostly in the Bulletin du Jardin botanique national de Belgique. Heinemann actively serached collaboration from well known European mycologists (Romagnesi, Dissing and Lange, Pegler, Horak, ...), translating, if necessary, their contributions into French. The flora underwent another change in incorporating collections from other African tropical regions, trying to be complete for central Africa. Microscopic details were drawn at the sme magnification (permetting easy comparison), and the drawings' legends coded, permitting their easy interpretation for non-native French readers. From frascicle 10 to fascicle 17, the last published in 1997, the keys were also translated in English, to broaden the use of the flora.

During the last three decades, the collectors and African collections in the herbarium of the National botanic garden of Belgium (BR) significantly increased in number and geographic origin. Mycology is also standing up in African countries as clearly exemplified by the number of African mycologists present at the AETFAT congresses in Meise (Belgium) (2000) and Yaoundé (Cameroon) (2007). The need is now felt for a series treating the mushrooms of Africa south of the Sahara. Out of the brainstorming at the National botanic garden of Belgium came the initiative to replace the Flore illustrée des champignons d'Afrique centrale in a Fungus Flora of Tropical Africa, building on the experiences with the former series. In oder to actively collaborate in this project we strongly encourage you to submit, in English or French, taxonomic revisions of not yet published fungal groups. We are also very pleased to announce that thanks to the Andrew W. Mellon Foundation (New York) funding of the African Plants Initiative Project all photographs and published iconographical material of the Meise collections will be avaialble soon on the Internet. With no doubt this tool will help to improve our knowledge of the African mycodiversity.

 

Le Jardin botanique national de Belgique possède une longue tradition dans l’étude de la mycoflore d’Afrique centrale. Les plus anciennes collections de champignons d’Afrique centrale conservées dans son Herbier sont plus que centenaires (1891, 1895), soit lorsque le Congo était encore la propriété privée de Léopold II, Roi des Belges. La publication la plus ancienne concernant la mycoflore congolaise est la Reliquiae Dewevreanae (1901) par De Wildeman & Durand.

Dès 1923, Mme M. Goossens-Fontana, professeur de dessin vivant au Jardin botanique d’Eala (Bas-Congo), s’adonna à l’aquarelle pour la représentation des champignons. De retour en Belgique, un mycologue amateur, M. Beeli, la motiva à récolter, à réaliser des exsiccata et des sporées et lui enseigna la microscopie des champignons. Elle séjourna ensuite à Binga et à Panzi (Kivu). Son herbier comprend 1870 numéros et 1331 aquarelles, principalement de champignons charnus et constitue un trésor exceptionnel. Dès 1926, Beeli, parfois avec Mme M. Goossens-Fontana, publia ses découvertes (environ 300 nouvelles espèces) dans la série Fungi Goossensiani.

En 1935, Beeli parvint à convaincre le directeur du Jardin botanique de Bruxelles, W. Robyns, à initier la publication d’une Flore illustrée des champignons du Congo, la Flore iconographique des champignons du Congo dont 17 fascicules ont été publiés jusqu’en 1970.

En 1972, P. Heinemann, le nouvel éditeur, présenta un nouveau concept pour la Flore, rebaptisée Flore illustrée des champignons d’Afrique centrale. Les descriptions étaient plus complètes, enrichies d’illustrations des caractères microscopiques. Les diagnoses étaient encore publiées séparément, généralement dans le Bulletin du Jardin botanique national de Belgique. Heinemann  bénéficiait de la collaboration de mycologues européens célèbres (Romagnesi, Dissing et Lange, Pegler, Horak, …) et traduisait, si nécessaire, leurs contributions en français. La Flore fut également l’objet d’un autre changement, en y incluant des collections d’autres régions d’Afrique tropicale tout en tâchant d’être complète pour l’Afrique centrale. Les détails microscopiques furent représentés au même grossissement de manière à permettre une comparaison aisée et les légendes des dessins codées afin de pouvoir être interprétées par des lecteurs non francophones. Du fascicule 10 au fascicule 17, le dernier publié en 1997, les clés furent également traduites en anglais afin d’élargir le public des utilisateurs de la Flore.

Durant les trois dernières décennies, les récolteurs et les collections africaines de l'Herbier du Jardin botanique national de Belgique (BR) augmentèrent significativement et leur origine se diversifia. La mycologie se développe également dans les pays africains comme en atteste le nombre de mycologues africains présents lors des congrès de l'AETFAT de Meise (2000) et de Yaoundé (2007). La nécessité d'une flore des champignons de l'Afrique sub-saharienne se fait sentir. De la réflexion que nous avons menée au Jardin botanque national de Belgique est née l'initiative de remplacer la Flore illustrée des champignons d'Afrique centrale par la Fungus Flora of Tropical Africa, enrichie de l'expérience des séries précédentes. Pour collaborer activement à ce projet, nous vous encourageons vivement à soumettre, en français ou en anglais, des révision taxonomiques de groupes non encore publiés. Nous sommes aussi très heureux d'annoncer que, grâce au financement du projet API par la Fondation Mellon, toutes les photographies et le matériel iconographique des collections de Meise seront bientôt disponibles sur Internet. Cet outil aidera sans aucun doute à améliorer notre connaissance de la mycodiversité africaine.

 

Acknowledgements

 

The author wishes to thank Jan Rammeloo, Director of the National Botanic Garden, Meise (Belgium) for the suggestion to work on this project and for his help. The author´s thanks also belong to the curators of the herbaria BR, BRNM, E, GENT, K, PC, PRM, WU, and ZT for the loan of type and non-type herbarium specimens and to B. Buyck (Paris, France), G. Eyssartier (Paris, France), I. Krisai-Greilhuber (Vienna, Austria), D.C. Mossebo (Yaoundé, Cameroon), P.J. Roberts (Kew, England), C. Sharp (Zimbabwe), A. Verbeken (Gent, Belgium), R. Watling (Edinburg, Scotland) for kindly submitting their collections. My thanks are due also to Zdeněk Pouzar (Prague, Czech Republic) for valuable nomenclatorical and taxonomic notes and Jan W. Jongepier (Veselí nad Moravou, Czech Republic) for linguistic improvement of this manuscript.

I am also very obliged to the "Fondation pour favoriser les recherches scientifiques en Afrique", which enabled my field trip to Benin in 1997, and also to Mr. André De Groote (Belgium) for logistics and help during my stay there.

The taxonomical study of this group as well as my collecting excursion to Cameroon in 2001 was supported by the Grant Agency of the Czech Republic (No. 206/01/0093).

 

Introduction

 

This monograph presents the first part of a taxonomic study of marasmioid genera in tropical Africa. It contains monographs of the very close genera Marasmius Fr. s. str., Gloiocephala Massee and Palaeocaephala Singer, and of the genus Setulipes Antonín, which forms a transient group to the genus Marasmiellus Murrill. The monograph of the latter genus will be the author´s next target.

In this book, 7 species of the genus Gloiocephala, 110 taxa (102 species and 8 varieties) of the genus Marasmius, 1 species of Palaeocephala, and 7 species of Setulipes are included. However, the approximate number of Marasmius species may be 2–3 times higher.

 

Material and methods

 

The region studied included the African continent between the tropics of Capricorn and Cancer except for the island of Madagascar. This island has a very different flora which does not belong to the tropical African one. The aim of this monograph is to summarise the knowledge about tropical African species of Marasmius s. str. and some related genera, and to stimulate and support studies of this group of fungi in the future.

The results of this monograph are based on studies of the author’s own collections from Benin and Cameroon and of herbarium specimens from the following herbaria: BR, BRNM, E, GENT, K, PC, PRM, S, UPS, WU, ZT, herb. D.C. Mossebo and material sent by B. Buyck, C. Douanla-Meli, G. Eyssartier, I. Krisai-Greilhuber, P.J. Roberts, C. Sharp, A. Verbeken and R. Watling. In total, about 850 specimens were studied.

Microscopic features are described from material mounted in Melzer's reagent, Congo Red, Cresyl Blue and ca. 5 % KOH. For macroscopic studies, an Olympus BX 50 light microscope was used.

 

Characters used for the identification of marasmioid and collybioid taxa (modified chapter from Antonín & Noordeloos 1993)

 

1  Macroscopical characters

1.1  Basidiocarps

Habit. Within the marasmioid genera several habit types are found: marasmioid, collybioid, omphalioid and pleurotoid. The collybioid habit is characterised by a neither umbilicate nor conical pileus, free or adnate lamellae, a tough context and the fact that context of pileus continuous with context of stipe, the marasmioid one by a plicate pileus, a horse-hair stipe and revivescent carpophores, the omphalioid one by a plano-convex to deeply unfundibuliform pileus and decurrent lamellae, and the pleurotoid one by the absent or lateral stipe. The first three are centrally stipitate.

Habit characters are not very important on generic level, with the exception of some pleurotoid genera, but at the infrageneric level (subgenus, section) habit characters are often used.

Reviving basidiocarps. Typical of many species of Marasmius and related genera, like Setulipes and Crinipellis, is the ability of the basidiocarps to revive after complete dehydratation, and to start the production of spores again. Huijsman (1971) came to the conclusion that not only Marasmius s. str. shows this reviving ability, but also species now classified to Micromphale, Marasmiellus, the 'peronatus'-group of Gymnopus, and members of some other genera of white- and brown-spored agarics.

 

1.2  Pileus

Shape. The shape of the pileus varies in the species, and also during the development of the basidiocarps. Many species have a hemispherical pileus when young that expands with age through convex to applanate. The centre can be obtuse, papillate, umbonate or depressed, but deeply umbilicate or infundibuliform pilei are very rare.

Colour. The colour in most species ranges from white to cream, brown, grey, violaceous, red, orange, ferrugineous, purple or almost black; the colour is sometimes very bright. In some species, the pileus may be striped, especially in sect. Sicci and Globulares.

Surface. Many species have a smooth pileal surface. However, taxa with diverticulate elements in the pileipellis often have a minutely tomentose pileus when observed under a strong lens. Some taxa have thin- to thick-walled hairs or setae. Species with a marasmioid habit have mostly a radially sulcate surface.

 

1.3  Lamellae

Development. Most taxa have well-developed lamellae that reach the margin of the pileus. In Marasmius, Gloiocephala, Palaeocephala and some other genera, however, species are known with lamellae which are reduced or vein-like, often not reaching the margin of the pileus. These vein-like structures are frequently furcate and/or anastomosing, or form an irregular veined pattern. Rarely, the lamellae are absent in young specimens and develop into a rugulose or veined hymenophore in mature specimens.

Attachment. The lamellae are free, adnexed, adnate or slightly decurrent. In some species, especially in sect. Marasmius, the lamellae are attached to a free collarium. In other sections, adnate lamellae partly sometimes become free from the stipe, tearing off part of the stipe-cortex, forming a collarium-like structure which is called a pseudocollarium or a false collarium.

Spacing. Lamellae can be very crowded to very distant. This feature, combined with the number of lamellae, and the presence or absence of lamellulae, forms an important distinguishing character.

Colour. Lamellae are usually white to pale cream-coloured, but in some species yellow-brown, ochraceous, grey-brown, brown, grey or reddish lamellae are found. The lamellar edge is usually concolorous with the sides, except for some species in sect. Sicci, where coloured edges occur. However, the colour of the lamella edge appears to be rather variable from one basidiocarp to another in one population, and therefore of limited taxonomic value.

 

1.4  Stipe

General characteristics. Most species have a centrally inserted, well-developed stipe. A pseudostipe occurs only in Chaetocalathus, Gloiocephala and Marasmiellus sect. Marasmiellus.

Consistency. Three main types of stipe are distinguished:

· filiform - very long and thin, like a horse-hair, usually considerably less than 1.5 mm thick.

· tough-cartilaginous - rather rigid and firm, more or less horny, not easy to break into pieces, usually relatively thin, ranging from 1–5 mm.

· fleshy - fibrous, but easily snapping across, and 2 to more than 15 mm thick.

Insertion. Basically, two types of insertion on or in the substrate are encountered:

·  insititious - attached directly to the substrate without special structures (typically in Marasmius sect. Marasmius).

· non-insititious - forming basal mycelium varying from a small basal disc to an abundant tomentum, spreading over or in the substrate, or very rarely rooting with a parts immersed in litter.

This is a character which plays an important role in the generic and infrageneric classification.

Surface. The surface of the stipe can be smooth and polished, as in many typically marasmioid species with filiform stipe, or be pruinose to densely tomentose. Some taxa have long, thin- or thick-walled to setiform hairs.

 

1.5  Rhizomorphs and sterile stipes

Rhizomorphs and sterile stipes (telepods) occur mainly in Marasmius, rhizomorphs rarely occur also in Marasmiellus and Setulipes, and are of importance at the species level.

 

1.6  Smell and taste

Many species have a fairly characteristic smell of onion, garlic, rotten cabbage, sewage or bitter almonds. In many cases the smell is an easy diagnostic character in the field. Farinaceous smell and taste are rarely encountered in the group of genera concerned.

 

2  Microscopical characters

2.1  Structure of the pileipellis

The structure of the pileipellis (Fig. 1) is one of the main characters used to delimit genera and sections within the genera. Three main types occur in the marasmioid genera: a cutis, a trichoderm, and a hymeniderm. In Collybia and Rhodocollybia the simplest types of cutis occur, consisting of narrow, wide, more or less cylindrical hyphae, sometimes embedded in a gelatinous matrix (ixocutis). A special type of cutis is the so-called Dryophila-type of pileipellis, which is characteristic of Gymnopus sect. Levipedes. In this type, the terminal elements are irregularly broadened, lobed or branched, and form patterns like a jig-saw puzzle when seen from above in a scalp. In Crinipellis and Chaetocalathus, the pileipellis is a cutis with transitions to a trichoderm, composed of long, dextrinoid hairs. In many species of Marasmiellus, and in those of Setulipes, the pileipellis consists of repent and/or ascending, often inflated diverticulate or irregular elements, often with numerous warts and/or small finger-like projections (Rameales-structure). The hymeniderm found in the genus Marasmius consists of smooth, clavate to globose elements, or of so-called “broom-cells” of the Rotalis- or Siccus-type, frequently mixed with well-developed pileocystidia or thick-walled setae. Circumcystidia (pileocystidia largely confined to a band around the pileus margin) are present in some species (M. thwaitesii).

 

2.2  Spores

In all genera concerned the spores are smooth, thin-walled, non-amyloid, non-dextrinoid, and not metachromatic in Cresyl Blue, except for the species of the genus Rhodocollybia, and some species of the genus Marasmius, sect. Globulares and Sicci, in which a part of the spores have thick, dextrinoid and cyanophilous walls (usually crassospores, sclerospores), and Chaetocalathus and partly Crinipellis, with often dextrinoid spores. Size and shape are very variable, are often important diagnostic characters at the specific, infraspecific, and to a minor extent also at the sectional level. In tropical species, basidiospores easily collapse and even in some well-preserved specimens it is sometimes impossible to find any spore.

 

2.3  Characters of the hymenium

The hymenium of many marasmioid species mainly consists of fusiform to clavate basidioles. Mature basidia are usually scarce, and collapse quickly after having released their spores. Size and shape of basidia and basidioles are of minor importance for the taxonomy. Crassobasidia (sclerobasidia, Clémençon 2004) are rarely present in some species among normal basidia and may be (weakly) dextrinoid.

Cheilocystidia (Fig. 2) occur frequently, and are often characteristic of the species, but may also be of importance at the sectional level in Marasmius. There is a larger diversity in size and shape, ranging from rather simple cylindrical to clavate, lageniform, coralloid ones or various types of broom-cells. Pleurocystidia are rare and, if present, of minor taxonomic importance, except for the delimitation of taxa in Marasmius sect. Chordales, Globulares and Sicci. Setae occur occasionally.

 

2.4  Trama

Lamellar trama is regular or subregular, never bilateral (except for the primordia in some species). Tramal hyphae are sometimes (slightly) gelatinised. Trama is mostly composed only of thin-walled hyphae; (slightly) thick-walled (and then more distinctly dextrinoid) hyphae are present in some species.

 

2.5  Stipitipellis and caulocystidia

The structure of the stipitipellis is of important diagnostic value. In typical Marasmii, i.e. those with either a filiform or a horny stipe, the stipitipellis is a cutis composed of very tightly packed, cylindrical hyphae, often with slightly to distinctly thickened walls. Often the hyphae are angular in cross section. This structure is probably responsible for the tough nature of the stipe. In typical Gymnopus species, the stipitipellis is less compact, the hyphal walls are not or only slightly thickened, and not coalesced. They are often rounded circular in cross section.

The stipe vesture (Fig. 3), whether smooth, striate, grooved or covered with cystidia or hairs, is of great importance at the sectional and species level. In Gymnopus, the sections are mainly distinguished by this character, in combination with the structure of the pileipellis. Crinipellis is characterised by the presence of long, dextrinoid hairs on the stipe-surface. At the species level the shape and size of caulocystidia is important.

 

2.6  Gelatinous context

In some species of the genera Marasmius, Gloiocephala, Marasmiellus and Rhodocollybia, the hyphae of the pileus, lamellae and the stipe context are embedded in a gelatinous substance. However, a gelatinous substance in pileus and/or lamellae is found to a certain degree in all genera concerned.

 

2.7  Chemical reactions of hyphae

The reactions of hyphal elements with Melzer's reagent (dextrinoid or non-dextrinoid, never amyloid) and Cresyl Blue (sometimes metachromatic) form a useful tool to distinguish sections in Marasmius. The complete absence of amyloidity have only one exception represented by Marasmius rhododendrorum Kalamees from the Caucasus with distinctly amyloid setae on pileus and stipe (Antonín 1993).

 

3  Ecological characters

In the genera considered here, many species are rather host or substrate-specific, which is a valuable tool to recognise them in the field. Some species are found almost exclusively on Monocotyledonous families like Gramineae, Cyperaceae, or Juncaceae (Marasmius curreyi, M. nigrobrunneus), some other have a relationship to specific host species. However, this fact is more important in European or North American species because of (almost) always well determined host species. In tropical African species, the substrate is in most cases identified as “dead leaves”, “dead twigs” etc. without any exact species identification. Therefore, the host specificity of most species is unknown.

The host/substratum specificity is far more important than other ecological or sociological features. Few species are for example considered specific to certain vegetation-types.

 

Description of macroscopic characters

 

A good macroscopic description belongs among basic conditions for the successful species identification. What is necessary to note in collected carpophores?

In a pileus, its size and shape, colour and a character of its surface (smooth, striate, sulcate, etc.) and in lamellae, their number, colour (edge and sides), a character of an attachment to stipe and the presence or absence of lamellulae. A size and shape, colour and its covering (smooth, fibrillose, tomentose, etc.) are necessary to note in a stipe description. Also the presence or absence of basal structures (basal mycelium, hairs, etc.) represents a very important feature. The presence or absence of such structures belongs to basic characters for a delimitation of some genera (e. g. Marasmiellus x Gymnopus). A smell of carpophore context and sometimes also their taste may lead to the successful species identification.

For a colour description, the use of a colour chart is recommended. Among the most used ones belong e.g. those by Kornerup & Wanscher (“Methuen handbook of colour”), Maerz & Paul (“A dictionary of color”).

 

Abbreviations

 

The following abbreviations are used: E = quotient of length and width of the spores, Q = the mean value of E in all collections studied, L = number of entire lamellae, l = number of lamellulae between each pair of entire lamellae.

In the illustrations H ba stands for basidia, sp for basidiospores, H chcy for cheilocystidia, H plcy for pleurocystidia, H cy for hymenial cystidia; H se for hymenial setae, H pa for paraphysoid cells, P pe for pileipellis, P cy for pileocystidia, P se for pileosetae, S pe for stipitipellis, S cy for caulocystidia, S se for caulosetae and BM for basal mycelium. The scale bar always represents 20 μm.

Authors of fungal names are cited according to Kirk & Ansell (1992), colour abbreviations are according to Kornerup & Wanscher (1983), and herbarium abbreviations follow Holmgren (2003).

In the enumeration of revised specimens for each taxon, a question mark is used in case of doubtful determination.

 

Key to marasmioid and collybioid genera

 

1. Pileipellis hymeniform (even in maturity) ............................  2

1*. Pileipellis never hymeniform, or rarely hymeniform only in very young pilei ....................  4

 

2. Carpophores large to small; hymenophore well-developed, in some cases reduced; pileipellis of smooth or broom-cells; gloeocystidia never present ...................  Marasmius

2*. Carpophores always tiny to small; hymenophore reduced or with veins or scattered lamellae; pileipellis of smooth cells; gloeocystidia present or all cystidia absent .................... 3

 

3. Cystidia (at least gloeocystidia) present ......................  Gloiocephala

3*. Cystidia absent in all parts of the carpophore ..............  Palaeocephala

 

4. Stipe insititious or subinsititious ..............  5

4*. Stipe never insititious ................... 8

 

5. Pileus and sometimes also stipe with long, often thick-walled setoid hairs .................  6

5*. Long, thick-walled setoid hairs absent ......................................................  7

 

6. Carpophores marasmioid to collybioid; stipe central ................................. Crinipellis (not included here)

6*. Carpophores pleurotoid; stipe lateral ...............................................  Chaetocalathus (not included here)

 

7. Tramal hyphae never dextrinoid; stipe (sub)insititious ........................  Marasmiellus (not included here)

7*. Tramal hyphae, at least stipe medulla hyphae dextrinoid; stipe insititious or subinsititious .............. Setulipes

 

8. Spore print pink to ochraceous orange when fresh; basidiospores usually dextrinoid and cyanophilous; pileipellis a simple cutis or ixocutis ...................... Rhodocollybia (not included here)

8*. Spore print white to cream-coloured; spores neither dextrinoid nor cyanophilous; pileipellis of smooth or irregular or diverticulate hyphae ........................  9

 

9. Carpophores small, often growing from a sclerotium or mummified remnants of fungal carpophores; pileipellis a cutis or ixocutis consisting of narrow, cylindrical hyphae, without projections and not diverticulate ......  Collybia (not included here)

9*. Carpophores usually larger, growing on other substrata, only rarely from a sclerotium; pileipellis a cutis to trichoderm consisting of hyphae with few to numerous projections or with lobed to coralloid terminal elements ...... Gymnopus (not included here)

 

 MARASMIUS Fr.

 

Marasmius Fr., Fl. Scan.: 339 (1836) (nomen conservandum). Type species (fixed by conservation): Marasmius rotula (Scop.: Fr.) Fr.

Heliomyces Lév., Ann. Sci. Nat., sér. 3, 2: 117 (1844), lectotype: H. elegans Lév. (Donk 1962); Androsaceus Pat., Hyménomyc. Eur.: 105 (1887), lectotype: Agaricus rotula Scop.: Fr. (Donk 1962); Marasmius, subgen. Collybiopsis J. Schröt. in Cohn, Kryptog.-Fl. Schles. 3(1): 559 (1889), Collybiopsis (J. Schröt.) Earle, Bull. N. Y. Bot. Gard. 5: 415 (1909), lectotype: Agaricus calopus Pers.: Fr. (Donk 1962); Mycenitis Earle, Bull. N. Y. Bot. Gard. 5: 414 (1909), holotype: Marasmius alliaceus (Jacq.: Fr.) Fr.; Scorteus Earle, Bull. N. Y. Bot. Gard. 5: 415 (1909), holotype: Marasmius oreades (Bolton: Fr.) Fr.; Tephrophana Earle, Bull. N. Y. Bot. Gard. 5: 427 (1909), holotype: Collybia fimicola Earle; Polymarasmius Murrill, N. Amer. Fl. 9: 286 (1915), holotype: Marasmius multiceps Berk. & M.A. Curtis.

 

Basidiocarps marasmioid or collybioid, small to large, revivescent. Pileus membranaceous to moderately fleshy, white or pigmented, smooth, glabrous, grooved or deeply radially sulcate; lamellae well-devel­oped or slightly to strongly reduced, rarely lacking, sometimes attached to a free or adnexed, sometimes only a partially developed collarium; mostly white, cream or (ochraceous) yellow. Stipe central, eccentric or lateral, rarely lacking, insititious or non-insititious (with basal mycelium, sometimes rooting), filiform to fleshy, then often cartilaginous, smooth or finely grooved; glabrous, pruinose, pubescent, hirsute, furfuraceous or squamulose; rhizomorphs and sterile stipes present or absent. Smell none or distinct, then often strong, like onions, garlic or bitter almonds. Spore print white to pale cream.

 

Basidiospores ellipsoid, cylindrical, amygdaliform, lacrimoid, fusoid or clavate, hyaline, smooth, thin-walled, inamyloid, non-dextrinoid, acyanophilous. Basidia subcylindrical to clavate, 2- or 4-, rarely 1- or 3-spored, hyaline. Basidioles cylindrical, clavate, fusoid, hyaline, thin-walled. Cheilocystidia present or absent, often present in the form of broom-cells, and then usually similar to those found in pileipellis, sometimes clavate or irregularly lobed. Pleurocystidia present or absent, clavate, cylindrical, fusiform, lageniform or lecythiform, hyaline or with pale yellow walls, sometimes with refractive content. Hyphae of trama subregular or irregular, cylindrical or inflated, sometimes branched, thin-walled, hyaline or pigmented, clamped or clampless. Pileipellis hymeniform, composed of smooth elements or broom-cells of the Siccus- or Rotalis-type. Pileocystidia or pileosetae sometimes present. Stipitipellis a dry cutis composed of slightly to distinctly thick-walled, cylindrical hyphae, sometimes more or less trichodermal with patent, thin-walled hyphal projections. Well-developed caulocystidia or setae frequently present. Clamp-connections mostly present and abundant, rarely absent.

 

Notes. Saprophytic, sometimes parasitic and causing plant diseases, on living or dead parts of herbaceous and woody plants. Systematically, according to the “classical” system of fungi (Hawksworth & al. 1995), it belongs to family Tricholomataceae R. Heim ex Pouzar. However, according to a new system based mostly on molecular methods (Kirk & al. 2001), it rather belongs to an independent family Marasmiaceae Kühner. 

The large genus Marasmius Fr. is cosmopolitan, with the main distribution in tropical regions. It contains about 600 species world-wide, and over 1600 epithets have been published in the genus. It belongs to the most common and the most important genera in all tropical regions; its species are especially very common in tropical rain forests. Most species are saprophytic and play a very important role in the decomposition of leaf and woody litter and in the turnover of energy. Several species also parasitise on economically important plants. From this point of view, it is very important to have a monographic study of this genus as a basis for further studies.

 

Brief history of Marasmius collections from tropical Africa

 

A complete history of the marasmioid and collybioid genera was published by Antonín & Noordeloos (1993). 

In comparison with other tropical regions, Africa is somewhat late in the research of marasmioid fungi. Studies of South American Marasmius species were carried out especially by Dennis (1958, 1961, 1968), Dennis & Reid (1957) and Singer (1960, 1965b, 1969), a monographic study by Singer (1976); studies in south-east Asia especially by Corner (1994, 1996), Desjardin & al. (2000), Petch (1948) and Pegler (1986), and in Australia by e.g. Desjardin & Horak (1997), Desjardin & Petersen (1989a) and Desjardin, Wong & Hemmes (1992).

Among the first mycologists collecting and publishing their more or less numerous marasmioid collections from tropical Africa is e.g. Eichelbaum (1907). At the same time, P. Hennings published a series of papers (Hennings 1893, 1895a,b, 1897, 1898a,b, 1900, 1901, 1902, 1904, 1905a,b) describing a lot of new species based on fungi collected by others (e.g. Dusén, Eichelbaum, Engler, Preuss, Staudt, Zenker and Zimmermann). Unfortunately, most of his collections, especially types, were destroyed during World War II in Berlin. M. Beeli published a long series of papers (Beeli 1923a,b, 1926, 1927a,b,c, 1928a,b, 1929, 1930, 1931, 1932, 1933, 1936a,b, 1938, 1940) also based on collections of other people (e.g. Deighton, Ghesquière, Goossens-Fontana, Vanderyst, etc.). Most of his collections are preserved in the Herbarium of the National Botanic Garden in Meise, Belgium (BR). Several Marasmius species were also described by Patouillard (1924, 1928, Patouillard & Hariot 1893; his collections are preserved in the Herbarium of the National Museum in Paris, France, PC), and de Seynes (1897; some collections are in the herbarium BR).

The last larger studies of the genus Marasmius from tropical Africa were published 40 years ago (Singer 1964, 1965a). However, these studies were based on limited number of specimens preserved in one herbarium (BR). These studies included 50 Marasmius species. Since that time no real monographic study has appeared. Descriptions or data of some species can be found scattered in other African literature, e.g. Dade (1940), Deighton (1936), Heim (1930, 1948), Hendrickx (1948), Morris (1990), Nicholson (1989), Pegler (1966, 1967, 1968, 1975, 1982), Pegler & Rayner (1969), Williamson (1976), Zoberi (1972). Pegler (1977) published a preliminary flora of agarics based on his collections from East Africa, which contains 43 Marasmius species.

 

Key to sections of the genus Marasmius

 

1. Pileipellis hymeniform ........................................  2

1*. Pileipellis not hymeniform but a cutis, with the Ramealis-structure or irregularly arranged broom-cells ..... sect. Fusicystides

 

2.   Lamellae attached to a distinct collarium, stipe always insititious .................................  sect. Marasmius

2*. Lamellae not attached to a distinct collarium, sometimes a pseudocollarium present, but in this case stipe not insititious ...  3

 

3. Pileipellis composed of broom-cells or cells with numerous digitate projections ...................................  4

3*. Pileipellis composed of smooth cells and/or a mixture of smooth and broom-cells ..............................  12

 

4. Pileus white, stipe always central, pubescent, insititious; pileipellis cells with digitate projections (not true broom-cells) ............... sect. Epiphylli, subsect. Epiphylloidei

4*. Pileus pigmented, stipe central to lateral, insititious or not insititious; pileipellis of true broom-cells .........  5

 

5. Pileipellis composed of broom-cells of the Rotalis-type; stipe insititious; trama hyphae non-dextrinoid ...... sect. Hygrometrici

5*. Pileipellis composed of broom-cells of the Siccus-type ............................  6

 

6. Stipe eccentric or rudimentary ............................  sect. Neosessiles

6*. Stipe always central or slightly eccentric .....................................  7

 

7. Stipe insititious; hyphae non-dextrinoid .................................. sect. Leveilleani

7*. Stipe not insititious; hyphae dextrinoid or non-dextrinoid ..........................  8

 

8.  Hyphae dextrinoid (sect. Sicci) ................................ 9

8*. Hyphae non-dextrinoid ..........................  sect. Inaequales

 

9.  Setae present on pileus or stipe .............  sect. Sicci, ser. Spinulosi

9*. Setae absent ........................... 10

 

10. Caulocystidia present .......................  sect. Sicci, ser. Atrorubentes

10*. Caulocystidia absent (except for the stipe apex where broom-cells may be present) ............ 11

 

11. Pleurocystidia present .....................  sect. Sicci, ser. Haematocephali

11*. Pleurocystidia absent ..........................  sect. Sicci, ser. Leonini

 

12. Carpophores small; pileus white to white-off; stipe insititious, filiform; lamellae vein-like to well-developed ..... sect. Epiphylli

12*. Carpophores larger; pileus usually pigmented; stipe not insititious; lamellae always well-developed .... 13

 

13. Context hyphae non-dextrinoid ..........................  sect. Chordales

13. Context hyphae dextrinoid .............................  14

 

14. Thick-walled setae present on pileus and/or stipe ..................  sect. Sicci, ser. Spinulosi

14*. Thick-walled setae absent ............................  sect. Globulares

 

Sect. Marasmius

 

Marasmius, II. Mycena, 2. Rotulae Fr., Epicr.: 384 (1838); Marasmius B. Rotulae Quél., Enchir.: 145 (1886); Marasmius sect. Rotulae Kühner, Botaniste 25: 98 (1933).

Marasmius, I. Rotularia J. Schröt. in Cohn, Kryptog.-Fl. Schles. 3(1): 556 (1889).

Marasmius, “sect.  Setipedes, α Collariati Bataille, Marasmes Eur.: 26 (1919).

Marasmius sect. Pararotulae Singer, Sydowia 18: 140 (1965).

Type species: Marasmius rotula (Scop.: Fr.) Fr.

 

Basidiocarps small, marasmioid. Pileus usually less than 20 mm broad, membranaceous, hemispherical to obtusely convex, mostly umbilicate, often with small papilla in umbilicus, radially grooved to sulcate, white, beige, grey, grey-brown, brown, yellow-brown, cinnamon-red to red-brown, often with distinctly darker centre. Lamellae well-developed, distant, without lamellulae or rarely with 1(–2) lamellulae, with well-developed, free, rarely partly attached collarium. Stipe insititious, filiform, rarely branched, usually whitish above, red-brown to black-brown below, shining; rhizomorphs or sterile stipes often present.

 

Basidiospores small to large, ellipsoid to slightly amygdaliform. Basidia 4- or 2-spored. Basidioles often fusoid, not exceeding the hymenium. Cheilocystidia present, in the form of broom-cells; pleurocystidia absent. Pileipellis hymeniform composed of broom-cells of the Rotalis- or Siccus-type. Stipitipellis a cutis composed of compact, smooth hyphae. Clamp-connections present. Hyphae of context usually dextrinoid, at least in stipe, sometimes non-dextrinoid.

 

Notes. Species of this section are frequent in the tropics. Taxonomically, this section is very complicated. Differential features of species are based mostly on macroscopic characters and size and shape of basidiospores.

In comparison with studies by Singer (1964, 1965a), I excluded two species from this section. Marasmius beelianus Singer (= M. epiphyllus var. congolensis Beeli) was mentioned by Singer (1964, 1965a) as belonging to this section. However, Beeli (1928a) did not describe the lamellae as collariate (“lamelles adnées-décurrentes …”) and neither is there a collarium in the type specimen. Moreover, the presence of up to 37 μm long pleurocystidia justifies its place in sect. Sicci (for details see there).

Singer (1964, 1965a) published a collection by J. Louis 3551 (BR 11513-67) as Marasmius subrhodocephalus Henn. In those carpophores, studied by Singer, however, the collarium was not developed! This is quite clear also on dry specimens (two carpophores). Moreover, it seems that its stipe in not insititious. Therefore, M. subrhodocephalus Henn. s. Singer (1964, 1965a) belongs to sect. Sicci, however, it is not possible to exactly identify it. Because of the absence of the type-specimen of M. subrhodocephalus Henn., I consider it a dubious name (see also chapter “Excluded and dubious names”).

 

 Key to tropical African species

 

1.  Pileipellis with broom-cells of the Rotalis-type (subsect. Marasmius) ........................  2

1*. Pileipellis with broom-cells of the Siccus-type (subsect. Sicciformes) ........................  8

 

2. Pileus cinnamomeous red; lamellae moderately close, L = 18–20; basidiospores rather large, 8.5–12.5 x 3.7–5.4 μm ..................................  1. M. louisii

2*. Pileus differently coloured, never red or reddish; lamellae more distant; basidiospores smaller ............. 3

 

3. Pileus very small, 0.5–1.5 mm broad; lamellae very distant, L = 7–8(–9); stipe very short (1.2–4 mm long), arising directly from long and often branched rhizomorphs; pileipellis cells of both Rotalis- and Siccus-types ............. 2. M. cupressiformis

3*. Pileus broader; lamellae more numerous; stipe longer and arising from substratum; pileipellis consisting of only one type of cells ...........................................  4

 

4.  Pileus white or whitish to cream when fresh, then pale beige; lamellae distant, L = 11–17 .................... 5

4*. Pileus differently coloured OR lamellae more distant (L < 12) ................................  6

 

5.   Stipe up to (7–)15–30(–80) mm long; basidiospores 6.5–10.0 x 3.5–4.7 μm ..............   3.1. M. rotalis var. rotalis

5*.Stipe up to 140 mm long; basidiospores 7.7–10.1(–10.8) x 4.6–5.4 μm ....  3.2. M. rotalis var. latisporus

 

6.  Lamellae 9–12 .......................................  4. M. apatelius

6*. Lamellae 10–16 ............................................ 7

 

7. Pileus brown, without central papilla; lamellae with pale grey-brown edges; walls of stipitipellis hyphae olivaceous green in KOH ...................................................  6. M. colorimarginatus

7*. Pileus light brown to brownish orange; lamellae with concolorous edges; walls of stipitipellis hyphae brown or grey-brown, never olivaceous green in KOH .........................  5. M. somalomoensis

 

8. Stipe short (up to 10 mm long), arising directly from long and often branched rhizomorphs ................. 9

8*. Stipe distinctly longer and arising from substratum or from both substratum and rhizomorphs ........... 10

 

9.   Pileus whitish, then pale beige brownish with dark chestnut brown centre; stipe very short (1.2–4 mm long); pileipellis cells of both Rotalis- and Siccus-types; basidiospores 6.2–8.9 x 3.5–5.0 μm ........................ 2. M. cupressiformis

9*. Pileus pale brown to brown, then often pale orange-brown or reddish brown, with very dark papilla; stipe slightly longer (3–10 mm); pileipellis cells only of the Siccus-type; basidiospores 7.7–12 x 4.2–6.2 μm ..................  9. M. crinisequi

 

10.  Basidiospores very large, 16–26 x 4.2–5.8 μm, clavate or narrowly lacrimoid ...... 11. M. yangambiensis

10*. Basidiospores always distinctly smaller, up to 18 μm long .........................  11

 

11. Pileus white, later pale yellow to pale yellow-orange, with a distinct black-brown or black central papilla; stipe straw coloured when young .......................................... 8. M. conicopapillatus

11*. Pileus distinctly coloured also when young, papilla (if present) concolorous or dark coloured; stipe black-brown also when young .......................................................................... 12

 

12. Basidiospores on average larger than 10 μm ................................ 13

12*. Basidiospores on average smaller than (or about) 10 μm ................................. 15

 

13. Basidiospores 13.5–18 x 4.5–7.5 μm; stipe orange to deep orange ................... 12. M. aurantiostipitatus

13*. Basidiospores smaller, up to 13.5(–14.5) μm long and up to 6.0 μm broad; stipe black or black-brown .................... 14

 

14. Lamellae 6–10; basidiospores (3.0–)3.5–4.5(–5.0) μm wide, lacrimoid, subfusoid; basidia 20–26 μm long; growing on dead leaves .......................................................... 13. M. guyanensis

14*. Lamellae 8–12; basidiospores 4.5–6.0 μm wide, fusoid; basidia 25–32 long; growing on dead twigs ...... 14. M. lovedalensis

 

15. Pileus grey, grey-brown or fuligineous brown; stipe very long, 13–73 x 0.1–0.5 mm; pileipellis cells with dark brown walls in KOH .......................................... 15. M. nigrobrunneus

15*. Pileus never with grey tinge, but red-brown, orange-brown, reddish; stipe distinctly shorter (up to 38 mm); pileipellis cells never with dark brown walls in KOH .................................. 16

 

16. Lamellae very distant (L = 6–9); stipe 4–12 mm long; basidiospores (8.0–)9.0–10.5(–12) x (4.5–)5.0–6.0 μm; cheilocystidia 10–23 x 6.5–12 μm large; growing on dead grass remnants ...................................... 10. M. curreyi var. distantifolius

16*. Lamellae more numerous (L = 8–14); stipe 7–38 mm long; basidiospores 7.7–9.6(–10) x (3.8–)4.2–5.4 μm; cheilocystidia 9.5–14.5 x 5.4–7.7 μm large; growing on dead leaves ........................ 7. M. subruforotula

 

Species descriptions

 

Subsect. Marasmius

 

Marasmius sect. Pararotulae Singer, Sydowia 18: 336 (1965); Marasmius sect. Marasmius, subsect. Pararotulae (Singer) Singer, Fl. Neotropica Monogr. 17: 92 (1976). Type species: M. pararotula Singer.

Marasmius sect. Marasmius, subsect. Penicillati Singer, Fl. Neotropica Monogr. 17: 121 (1976). Type species: M. graminum (Lib.) Berk. & Broome.

Type species: Marasmius rotula (Scop.: Fr.) Fr.

 

Note. Pileipellis consisting of broom-cells of the Rotalis-type.

 

1. Marasmius louisii Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 332 (1964). – Type: Democratic Republic of Congo, Yangambi, Isalowe Nature Reserve, 22 May 1939, J. Louis 14931 (BR 11487–41, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 332–334 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 258 (1965).

 

Pileus 2–3 mm broad, distinctly broader than high, convex, umbilicate, with small papilla in umbilicus, slightly tomentose, striate, cinnamomeous red, between “cochin” and “kis kilim” (Maerz & Paul) when dry, uniformly coloured or with grey or fuligineous papilla. Lamellae moderately close, L = 18–20, l = 0, collariate, broad (1 mm), white, with concolorous edge. Stipe 10–13 x 0.1–0.2 mm, filiform, insititious, smooth and glabrous, black-brown to black. Rhizomorphs absent. (According to Singer 1964, 1965a).

 

Basidiospores 8.5–12.5 x 3.7–5.4 μm, E = 1.8–2.7, Q = 2.2, ellipsoid, smooth, thin-walled, hyaline. Basidia 17–20 x 6.9–8.5 μm, 4-spored, clavate. Basidioles 19–27 x 6.9–9.2 μm, clavate, cylindrical or fusoid. Cheilocystidia similar to pileipellis cells, 15.5–19.5 x 6.9–7.7 μm, clavate, diverticulate in upper part. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 12–18 x 6.9–11.5 μm, clavate to subcylindrical, thin- to slightly thick-walled, with yellow-brown walls in KOH; projections wart-like to cylindrical, slightly thick-walled, obtuse, up to 5.0 x 1.0 μm. Pileocystidia absent. Stipitipellis a cutis composed of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae with yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 4

 

Chemical reactions. Stipe medulla and cortex hyphae weakly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves in a tropical forest.

 

Distribution. Known only from the type locality in the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Yangambi, Isalowe Nature Reserve, 22 May 1939, J. Louis 14931 (BR 11487–41, holotype).

 

Notes. Marasmius louisii is characterised by having a cinnamomeous red pileus, moderately close lamellae and rather large basidiospores. The pileus colour of this species is rather unique for species of subsect. Marasmius – it is more common in subsect. Sicciformes. Marasmius buzae Dennis, from Bolivia and Venezuela, has an orange-ferrugineous or orange-ochraceous pileus, less numerous lamellae (L = 7–11) and broader basidiospores, 8.5–13 x (5–)5.7–9(–10.3) μm; M. nebularum Singer, from Colombia, also has a dark  ferrugineous pileus but its basidiospores are smaller (6.2–7.5 x 3.2–3.7 μm); M. aequatorialis Singer, from Ecuador, with a brightly rusty orange or orange coloured pileus also has smaller basidiospores (5.5–6.5 x 4–4.5 μm) (Singer 1976).

 

2. Marasmius cupressiformis Berk.

Berkeley, J. Bot. (Hooker) 8: 140 (1856). Type: Brazil, Amazonas, Panuré, on leaf, Spruce 75, ex herb. M.J. Berkeley, No. 75 (K(M) 99701, holotype).

 

Selected descriptions and icons. Nicholson, Nigerian Field 61: 138 (1996); Singer, Bull. Jard. Bot. Etat Brux. 34: 329–330 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 257 (1965).

 

Pileus 0.5–1.5 mm broad, almost conical with prominent papilla, then convex to umbilicate, slightly depressed and umbonate in umbilicus, slightly tomentose, sulcate, whitish, then (always in herbarium specimens) pale beige brownish with dark chestnut brown centre. Lamellae distant, L = 7–8 (–9), l = 0, collariate, broad, white, often concolorous with pileus in herbarium specimens. Stipe 1.2–4 x 0.1 mm, filiform, smooth, glabrous, insititious, attached directly to rhizomorphs, black or chestnut brown-black. Context almost absent. Rhizomorphs black, long, often branched, wider than stipe (0.2–0.3 in diam.), smooth and glabrous. (According to Singer 1964, 1965a)

 

Basidiospores 6.2–8.9 x 3.5–5.0 μm, E = 1.5–2.0, Q = 1.8, ellipsoid or sublacrimoid, thin-walled, smooth. Basidia 19–20 x 5.5–7.0 μm, 4-spored, clavate. Basidioles 11.5–23 x 2.7–7.0 μm, clavate, cylindrical, subfusoid. Cheilocystidia shaped as broom-cells of both the Rotalis- and Siccus-types, 9.2–17 x 5.8–6.9 μm, clavate, diverticulate in upper part, similar to pileipellis cells. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, branched, up to 8.0 μm wide. Pileipellis a hymeniderm composed of both forms of broom-cells – Rotalis- and Siccus-type, 10.0–19.5 x 6.2–10.0 μm, clavate to subcylindrical, entirely thin- to slightly thick-walled above, with hyaline to pale (greyish) yellow walls in KOH; mixed with numerous, 13–21.5 x 9.2–12.5 μm, irregular, lobate to coralloid, thick-walled cells, sometimes with several projections, with distinctly yellow-brown walls in KOH. Pileocystidia absent. Stipitipellis a cutis composed of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae with yellow-brown walls in KOH. Caulocystidia absent; adpressed to suberect, cylindrical, terminal cells sometimes present. Clamp-connections present in all tissues. — Fig. 5

 

Chemical reactions. Stipitipellis hyphae weakly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead wood.

 

Distribution. Known only from the Democratic Republic of Congo and Brazil.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. unknown locality, 1923, J. Ghesquière 1355 (BR 11435–86).

Nigeria. Cross River State, Uyo-Calabar Road, Okpokong Bridge, 15 Sept. 1990,  R.A. Nicholson 740 (K(M) 23035).

 

Revised specimens from other regions.

Brazil. Amazonas, Panuré (= São Jerônimo), on leaf,  Spruce 75, ex herb. M.J. Berkeley, No. 75 (K(M) 99701, holotype).

 

Notes. Marasmius cupressiformis is characterised by having a very small, whitish, then pale beige brownish pileus with a dark chestnut brown centre, very distant lamellae, a very short stipe arising directly from long and often branched rhizomorphs and by the presence of pileipellis cells and cheilocystidia of both Rotalis- and Siccus-types. However, the ratio between both types is different in various collections. In the type collection, the Siccus-type cells are dominating.

Marasmius multiceps Berk. & M.A. Curtis, from Cuba and Guatemala, differs by more numerous lamellae (L = 9–16) and a larger, 1.5–6 mm broad pileus; however, it has only Rotalis-type broom-cells in the pileipellis (Singer 1976); M. pallenticeps Singer, known from Argentina and New Zealand, has less numerous lamellae (L = 5–7), a longer stipe (10–25 mm), and only Siccus-type broom-cells in the pileipellis (Desjardin & Horak 1997; Singer 1976).

 

3. Marasmius rotalis Berk. & Broome

Berkeley & Broome, Journ. Linn. Soc., Bot. 14: 40 (1873).

 

3.1. M. rotalis var. rotalis

Type: Sri Lanka, Central Province, Peradeniya, Jan. 1869, Thwaites 810 (K(M) 92650, holotype).

 

Selected descriptions and icons. Corner, Nova Hedwigia Beih. 111: 92 (1996); Desjardin & Horak, Bibl. Mycol. 168: 69–70 (1997); Desjardin & al., Sydowia 52: 111–113 (2000); Morris, Kirkia 13(2): 341 (1990); Pegler, Kew Bull. Addit. Ser. 6: 161–163 (1977); Pegler Kew Bull. Addit. Ser. 12: 149 (1986); Petch, Tr. Brit. Mycol. Soc. 31: 33 (1948); Singer, Bull. Jard. Bot. Etat Brux. 34: 331–332 (1964); Singer, Flore Icon. Champ. Congo 14: 257–258 (1965).

 

Pileus 1.5–9 mm broad, hemisphaerical when young, then convex to low convex, almost applanate in the end, umbilicate, with obtuse papilla at centre, low or with slightly uplifted margin when old, plicate-striate, white or whitish to cream when fresh, then pale beige (brownish, “Wigwam” in Maerz & Paul, when dried) with concolorous, but mostly grey-brown, brown-black or beige greyish papilla. Lamellae moderately distant, L = 10–17, l = 0(–1), broad, collariate, white to pale cream (3–5A2), with concolorous edge. Stipe (7–)15–30(–80) x 0.1–0.2 mm, filiform, glabrous, smooth, lustrous, insititious, fuligineous to fuligineous-black or brown-black, with apex concolorous with lamellae. Context thin. Rhizomorphs present, black, thin, glabrous. Pl. 1

 

Basidiospores 6.5–10.0 x 3.5–4.7 μm, E = 1.5–2.4, Q = 1.9–2.5, ellipsoid, subfusoid, sublacrimoid, thin-walled, hyaline. Basidia 21–26 x 7.5–9.0 μm, 4-spored, clavate. Basidioles 13–28 x 4.0–9.0 μm, cylindrical, clavate, fusoid. Cheilocystidia similar to pileipellis cells, 17–25 x 7.0–20 μm, clavate, thin-walled. Pleurocystidia absent. Trama hyphae cylindrical, ± thin-walled, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 12–28(–37) x 7.0–18(–25) μm, thin- or slightly thick-walled, (broadly) clavate; projections digitate, obtuse to subacute, slightly thick-walled, up to 4.0 x 1.0 μm. Stipitipellis a cutis composed of cylindrical, parallel, smooth, up to 5.0 μm wide hyphae. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 6

 

Chemical reactions. Pileus and lamellar trama hyphae non-dextrinoid, stipe medulla and cortex hyphae dextrinoid.

 

Ecology. Saprophytic, growing gregariously on dead leaves and twigs of dicotyledons in both savannah woodland and tropical forest.

 

Distribution. A pantropical species recorded in Africa (Benin, Cameroon, Democratic Republic of Congo, Kenya, Malawi, Nigeria, Tanzania, Uganda; Morris 1990, Singer 1964, 1965a, Pegler 1977), South America (Colombia; Singer 1976), Asia (Indonesia, Malaya, Sri Lanka; Corner 1996, Desjardin & al. 2000, Pegler 1986).

 

Revised specimens from tropical Africa.

Benin. Borgou Province, Wari Maro, 22 Aug. 1997, V. Antonín B97.82 (BR 101129–55); Atacora Province, Kota, 29 Aug. 1997,  V. Antonín B97.121 (BR 101165–91).

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 6 April 2001,  V. Antonín Cm 01.82 (BRNM 666076) ; Ibid., 8 April 2001,  V. Antonín Cm 01.49 (BRNM 666116); Ibid., 12 April 2001,  V. Antonín Cm 01.107 (BRNM 666156); South-West Province, Korup National Park, Mundemba, transect P22, Aug. 1990,  R. Watling (E); ? South-West Province, Korup National Park, transect P10, 22 March 1991,  R. Watling (E); ? Ibid., Mundemba, transect P24, 7 April 1990,  R. Watling (E).

Democratic Republic of Congo. Yangambi, Isalowe Reserve, 14 June 1939,  J. Louis 14934 (BR 11503–57).

Kenya. Central Province, Kiambu District, Muguga, 13 March 1968,  D.N. Pegler K 52 (K(M) 133687).

Malawi. Chifundi, Zomba Plateau, 26 Dec. 1981,  B. Morris 456 (K(M) 133686).

Nigeria. ? Cross River State, Calabar-Cameroon Road, 3 July 1990,  R.A. Nicholson 598 (K(M) 16773); ? Cross River State, Obudu Ranch, 25 April 1989,  R.A. Nicholson 190 (K(M) 7457); ? Cross River State, Uyo-Calabar Road, Itu Bridge, 1 July 1990,  R.A. Nicholson 563 (K(M) 18621).

Tanzania. West Usambara Mts., Mafi Hill, Kwalukonge stream, 27–28 Jan. 1985,  I. Krisai 3538 (WU).

Uganda. Buganda Province, Mengo District, Mawacota County, Mpanga Research Forest, 8 June 1968,  D.N. Pegler U 1246 (K(M) 133689); Ibid., 8 June 1968, D.N. Pegler U 1316 (K(M) 133690); Buganda Province, Mengo District, Mabira Forest, 9 June 1968,  D.N. Pegler U 1343 (K(M) 133691); Western Province, Bunyoro District, near Masindi, Budango Forest, 15 June 1968,  D.N. Pegler U 1488 (K(M) 133692).

 

Revised specimens from other regions.

Sri Lanka. Central Province, Peradeniya, Jan. 1869, Thwaites 810 (K(M) 92650, holotype).

 

Notes. Marasmius rotalis is characterised by having a rather small white or off-white pileus with a dark central papilla, rather close lamellae, a fuligineous to fuligineous-black or chestnut black stipe and by its growth on leaves of dicotyledons.

Descriptions of microscopic features in literature are identical with those described here, except for the slightly narrower basidiospores of the former - Desjardin & al. (2000): 7–9 x 3–4 μm; Desjardin & Horak (1997): 7.5–9 x 3–3.5 μm; Pegler (1977, 1986): 6–8.5 x 3–3.7 μm. However, my type revision showed that basidiospores may be up to 4.5 μm wide.

A similar species is M. leucorotalis Singer. It has more distant lamellae (L = 9–13) and smaller basidiospores (Desjardin & al. 2000; Pegler 1983; Singer 1976); M. rotuloides Dennis, from Trinidad, has smaller basidiospores (5–7.2 x 2–3.5 μm) (Dennis 1970; Singer 1976); M. pararotula Singer, from Bolivia, has broader basidiospores (7.5–8.3 x 4.8–5.5 μm); M. vergeliensis Singer, from Colombia, has a seldom elevated central white papilla, and M. cundinamarcae Singer, from Colombia, has more numerous lamellae (L = ± 20) and smaller basidiospores (5.5–7.2 x 2–3 μm) (all Singer 1976).

 

3.2. M. rotalis var. latisporus Antonín

Antonín, Mycotaxon 89(2): 427 (2004). Type: Burundi, Bururi Province, Bururi, 7 Feb. 1979,  J. Rammeloo 6577 (BR 11930–96, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 89(2): 427-428 (2004).

 

Pileus up to 4 mm broad, campanulate, umbilicate, with less distinct to distinct central papilla, distinctly sulcate, glabrous to minutely granulose, margin crenulate, pale beige, with or without black centre. Lamellae moderately distant, L = 14–17, l = 0(–1), collariate, white to whitish, with concolorous edge. Stipe very long, up to 140 x 1 mm, filiform, glabrous, smooth, lustrous, insititious, dark brown to black, apex concolorous with lamellae. Context thin. — Pl. 1

 

Basidiospores 7.7–10.1(–10.8) x 4.6–5.4 μm, E = 1.6–1.9, Q = 1.7, ellipsoid, ellipsoid-fusoid, thin-walled, hyaline. Basidia not found. Basidioles 15–27 x 5.0–10.0 μm, cylindrical, clavate, fusoid. Cheilocystidia similar to pileipellis cells, 15–29(–38) x 6.0–17 μm, clavate, thin-walled. Pleurocystidia absent. Trama hyphae cylindrical, ± thin-walled, up to 8 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 11–29 x 7.0–19 μm, thin- or slightly thick-walled, (broadly) clavate; projections digitate, obtuse, slightly thick-walled, up to 4.0 x 1.0 μm. Stipitipellis a cutis consisting of cylindrical, parallel, smooth, up to 5.0 μm wide hyphae. Caulocystidia absent. Clamp-connections present in all tissues. - Fig. 7

 

Chemical reactions. Stipe medulla and cortex hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. On dead fallen leaves.

 

Distribution. Known only from the type locality in Burundi.

 

Revised specimens from tropical Africa.

Burundi. Bururi Province, Bururi, 7 Feb. 1979,  J. Rammeloo 6577 (BR 11930–96, holotype).

 

Notes. Marasmius rotalis var. latisporus differs from the type variety in having a very long stipe and larger, especially broader basidiospores.

 

4. Marasmius apatelius Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 332 (1964). Type: Democratic Republic of Congo, Kisantu, 20 March 1907, H. Vanderyst s.n. (BR 11377–28, as M. friesianus, holotype).

 

Selected descriptions and icons. Pegler, Kew Bull. Addit. Ser. 6: 163. 1977. Singer, Bull. Jard. Bot. Etat Brux. 34: 332 (1964); Singer, Fl. Icon. Champ. Congo, 14: 258 (1965).

 

Pileus 1–6 mm broad, convex, umbilicate, with or without papilla in umbilicus, sulcate-striate, slightly tomentose, whitish (?) at first, then uniformly opaque and pale beige to brownish, up to flesh pink towards margin. Lamellae distant, L = 9–12, l = 0, collariate, sometimes furcate, broad, white. Stipe 10–25 x 0.1–0.2 mm, filiform, smooth, glabrous, insititious, fuligineous brown or black-brown, white or paler at apex. Context thin. Rhizomorphs only slightly developed, very thin. (According to Singer 1964, 1965a and Pegler 1977). — Pl. 1

 

Basidiospores 7.0–10.0 x 3.7–4.6 μm, E = 1.8–2.3, Q = 2.0, ellipsoid, thin-walled, smooth, hyaline. Basidia 16–20 x 4.0–5.0 μm, 4-spored, clavate. Basidioles 9.0–19 x 3.8–6.9 μm, clavate, fusoid, cylindrical. Cheilocystidia similar to pileipellis cells, 16–19.5 x 7.7–11.5 μm, clavate, diverticulate in upper part. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, branched, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 19–31 x 7.7–21 μm, clavate to (sub)vesiculose, thin- to slightly thick-walled; projections wart-like to digitate, slightly thick-walled, obtuse, up to 2.5 x 1.0 μm, with pale brownish-yellowish walls in KOH. Pileocystidia absent. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 5.0 μm wide hyphae with yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 8

 

Chemical reactions. Stipe medulla and cortex hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing gregarious on fallen dead leaves.

 

Distribution. So far known from the Democratic Republic of Congo, Tanzania and Uganda.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Kisantu, 20 March 1907, H. Vanderyst s.n. (BR 11377–28, as M. friesianus, holotype).

Tanzania. Tanga Province, Lushoto District, E. Usambara Mts., Amani, Bomole Hill, 20 April 1968, D.N. Pegler T 610 (K(M) 133699).

Uganda. Masaka District, Bukoko County, SW end of Kako forest, 5 May 1972, K. Arnstein Lye M 147 (K(M) 133700).

 

Notes. Marasmius apatelius is characterised by having a whitish, then uniformly opaque and pale beige to brownish pileus without a dark central papilla, distant lamellae and a ± brown stipe.

Very close is Marasmius leucorotalis Singer, which differs only in having a white to off-white pileus with a black conical central papilla. Marasmius rotuloides Dennis, from Trinidad, has smaller basidiospores (5–7.2 x 2–3.5 μm) (Dennis 1970; Singer 1976); M. manuripiensis Singer, from Bolivia, a paler (ochraceous to fuscous) central dot at the pileus centre, less numerous lamellae (L = 6–9) and larger basidiospores (7.5–11 x 4–5 μm) (Singer 1976); M. cundinamarcae Singer, from Colombia, has more numerous lamellae (L = ± 20) and smaller basidiospores (5.5–7.2 x 2–3 μm) (Singer 1976); M. rosulatus Desjardin & R.H. Petersen, from New Zealand, has a small pileus (0.2–0.5 mm) and less numerous lamellae (L = 3–5) (Desjardin & Petersen 1989a), and M. ubiquipallens Desjardin & E. Horak possesses a pale tan, slightly pale reddish orange tinged pileus and a stramineous to pale brown stipe (Desjardin & Horak 1997). 

 

5. Marasmius somalomoensis Antonín

Antonín, Mycotaxon 88: 66 (2003). Type: Cameroon, Sud Province, Somalomo, Dja Biosphere Reserve, 8 April 2001, V. Antonín Cm 01.42 (BRNM 666108, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 66–67 (2003).

 

Pileus 1–5 mm broad, convex, then low-convex, depressed at centre, with a small central papilla when young, absent when old, crenulate at margin, striate up to the centre, finely tomentose, light brown to brownish orange (6D4–5, 6C5), slightly darker, almost dark brown at central papilla, sometimes with white zone around papilla when dry. Lamellae moderately distant, L = 11–13, l = 0(–1), collariate, broadly adnate, rather broad, sometimes intervenose, whitish to pale (greyish) orange (4–5A2, 5B3), with concolorous edge. Stipe 10–17 x 0.1–0.2 mm, filiform, insititious, lustrous, twisted, concolorous with lamellae at apex, through dark brown to black brown towards base; sterile stipes present. Context membranaceous.

 

Basidiospores 7.0–10.0(–11) x 3.7–5.5 μm, E = 1.6–2.1, Q = 1.7–1.9, ellipsoid, ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidia 20–28 x 8.0–8.5 μm, 4- (rarely 2-)spored, clavate. Basidioles 15–28 x 4.0–10 μm, clavate, fusoid, cylindrical. Cheilocystidia shaped as broom-cells of the Rotalis-type, 14–30(–40) x 8.0–18(–22) μm, (broadly) clavate, thin-walled, hyaline, with slightly thick-walled, up to 3.0 x 1.5 μm projections; mixed with scattered similar smooth clavate cells. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, smooth to minutely incrusted, branched, up to 12 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 18–33 x 10–19 μm, entirely thin- to very slightly thick-walled, hyaline, dirty yellow to (ochraceous) brown in thick-walled parts, with up to 3.0 x 1.5 μm, slightly thick-walled warts with the same colour as the wall. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 6.0 μm wide hyphae, with brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 9

 

Chemical reactions. Stipe medulla and cortex hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves.

 

Distribution. Known only from Cameroon and the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 8 April 2001, V. Antonín Cm 01.42 (BRNM 666108, holotype); Ibid., 10 April 2001, V. Antonín Cm 01.79 (BRNM 666145); Ibid., 11 April 2001, V. Antonín Cm 01.91 (BRNM 666085).

Democratic Republic of Congo. Kivu, Irangi, 19 April 1972, J. Rammeloo Z 334 (GENT).

 

Notes. Marasmius somalomoensis is characterised by having a distinctly coloured pileus, moderately distant lamellae, rather large basidiospores and large basidia and basidioles.

In tropical Africa the closest species seems to be M. rotalis Berk. & Broome. It differs by the pileus being white or whitish to cream when fresh, then pale beige possessing a beige to grey-brown central papilla, more numerous lamellae (13–17), slightly narrower basidiospores (7.0–10.0 x 3.5–4.5 μm), smaller cheilocystidia (17–19 x 7.0–9.0 μm) and basidia (21 x 7.0 μm). Marasmius tubularis Petch, found in Sri Lanka, has a greyish brown pileus with a white margin and a black central papilla, less numerous lamellae (L = 8–12) with a coloured edge and smaller cheilocystidia (12–16 x 8–11 μm, Pegler 1986); M. plicatus Wakker, from Indonesia, has a pileus with an off-white umbilicus without a papilla, less numerous lamellae (L = 7–8) and slightly larger basidiospores (9–11 x 4–6 μm) (Desjardin & al. 2000); M. arimanus Dennis, from Trinidad, has a sepia or mummy brown pileus, less numerous lamellae (L = 7) and longer and narrower basidiospores (11–14 x 3.5–4 μm); M. dodecaphyllus Singer, from Bolivia, has a pileus with a flat, grey central dot, less numerous lamellae (L = 12), an umber brown stipe, and smaller basidiospores (7.5–8.2 x 3–4.5 μm); M. platyspermus Singer, from Argentina, has an at first whitish, then yellow beige pileus with a black central papilla, more numerous lamellae (L = 13–20) and grows on twigs and wood; M. tetrachrous Singer, from Bolivia, has an at first ferrugineous, then in various tinges rusty pileus with a black central papilla, less numerous lamellae (L = ± 9) and shorter and broader basidia (16.5–19.5 x 10–11 μm); M. tereticeps Singer, from Bolivia, has a cinnamon ochraceous pileus with a black central papilla, extraordinary broad lamellae and narrower basidiospores (8.2–11.2 x 3.5–4 μm); M. variabiliceps Singer var. variabiliceps, from Bolivia, has a pileus with a black central papilla, more numerous lamellae (L = ± 20) and a very long stipe (58–100 mm) (all Singer 1976).

 

6. Marasmius colorimarginatus Antonín

Antonín, Mycotaxon 88: 57 (2003). Type: Cameroon, Sud Province, Somalomo, Dja Biosphere Reserve, 9 April 2001, V. Antonín Cm 01.61 (BRNM 666128, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 57–58 (2003).

 

Pileus 8–10 mm broad, convex, with slightly inflexed crenulate margin, depressed at centre, without central papilla, plicate-striate, slightly tomentose, uniformly coloured, brown (7E5), only slightly darker at centre. Lamellae moderately distant, L = 13–14, l = 0(–1), collariate, broad, intervenose, cream (4A2), with pale grey-brown edge. Stipe 18–22 x 0.1–0.2 mm, filiform, insititious, smooth, glabrous, entirely black-brown. Rhizomorphs present. — Pl. 1

 

Basidiospores 8.0–10.0 x 3.5–5.5 μm, E = 1.7–2.3, Q = 2.0, ellipsoid to pip-shaped, thin-walled, hyaline, smooth. Basidia not found. Basidioles 15–28 x 4.0–8.5 μm, clavate, cylindrical, subfusoid. Cheilocystidia 16–31 x 9.0–15 μm, shaped as broom-cells of the Rotalis-type, (broadly) clavate, hyaline, thin-walled, with thin- to slightly thick-walled projections; scattered, diverticulate, hyphoid, prolongate cells present on lamellar edge. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin- to slightly thick-walled, branched, up to 15 μm wide, with hyaline to pale brownish walls (esp. in subpileipellis) in KOH. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 16–32 x 11–20 μm, (broadly) clavate, thin- to slightly thick-walled, with thick-walled projections and hyaline (thin-walled parts) or brown (thick-walled ones) walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 5.0 μm wide hyphae with distinctly olivaceous green walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 10

 

Chemical reactions. Stipe medulla and cortex hyphae slightly to distinctly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves in a tropical forest.

 

Distribution. Known only from Cameroon.

 

Revised specimens from tropical Africa.

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 9 April 2001, V. Antonín Cm 01.61 (BRNM 666128, holotype).

 

Notes. Marasmius colorimarginatus is characterised by a uniformly brown pileus without central papilla, intervenose lamellae with pale grey-brown edge, a stipe arising directly from substrate, rather broad basidiospores (8.0–10.0 x 3.5–5.5 μm) and stipitipellis hyphae with olivaceous green walls in KOH.

Among species with coloured lamella edge, Marasmius ubiquipallens Desjardin & E. Horak, from Papua New Guinea, has a pale tan, reddish orange tinged pileus and smaller basidiospores (7–9 x 4–4.5 μm) (Desjardin & Horak 1997); M. tubulatus Petch, from Sri Lanka, has a pallid, then greyish brown pileus with a white margin, a well-developed black papilla and smaller cheilocystidia (12–16 x 8–11 μm); M. buzae Dennis, from Bolivia and Venezuela, has an orange-ferrugineous or orange ochraceous pileus, more distant lamellae (L = 7–11) and broader basidiospores, 8.5–13 x (5–)5.7–9(–10.3) μm (Dennis 1970; Singer 1976); M. arimanus Dennis, from Trinidad, has a sepia or mummy brown pileus, less numerous lamellae (L = 7) and broader basidiospores, 11–14 x 3.5–4 μm; M. violeorotalis Singer, from Colombia, a violet-lilac pileus and narrower basidiospores (6.2–9.5 x 2.5–3 μm); M. vigintifolius Singer, from Bolivia and Brazil, has an ochraceous tan, brown or fuscous pileus, more numerous lamellae (L = 17–22) and a long stipe (14–56 mm); M. idroboi Singer, from Colombia, has a brown, paler radially striped pileus with a distinct central papilla, more numerous lamellae (L = 20–23) and smaller basidiospores (6.5–8 x 2.8–3.8 μm); M. baeocephalus Singer, from Ecuador, has a brown pileus with a white zone in the umbilicus and a well-developed dark papilla, less numerous lamellae (L = 9–11) and smaller basidiospores (6–8 x 3.5–4 μm). The South-American species M. variabiliceps Singer, which has several forms, differs by a rusty pileus with a black central papilla and larges basidiospores (10–11.7 x 4–4.8 μm) (var. mesites Singer), a deep brown to chestnut-ferrugineous pileus with a black central papilla and smaller basidiospores (7.5–8.8 x 4–4.8 μm) (var. tucumanensis Singer), an ochraceous pileus with a black central papilla and larger basidiospores (8.3–10.3 x 4.8–7 μm) (var. separatus Singer) or a ferrugineous, then reddish pileus with a flat black central papilla and less numerous lamellae, L = 10–13 (var. derubricans Singer). (all Singer 1976).

 

 

Subsect. Sicciformes Antonín

 

Marasmius sect. Marasmius, subsect. Sicciformes Antonín, Acta Mus. Moraviae, Sci. Nat., 76: 145 (1991).  Type species: Marasmius curreyi Berk. & Broome

 

Note. Pileipellis consisting of broom-cells of the Siccus-type.

 

7. Marasmius subruforotula Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 339 (1964). Type: Democratic Republic of Congo, Equateur Province, Eala, July 1907, L. Pynaert 1608 (BR 11515–69, holotype).

 

Selected descriptions and icons. Morris, Kirkia 13(2): 342 (1990); Pegler, Kew Bull. Addit. Ser. 6: 165–166 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 339–340 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 260–261 (1965).

 

Pileus 3–5 mm broad, convex to applanate, depressed or often umbilicate at centre, with distinctly acute, conical, lustrous central papilla, glabrous, sulcate-striate, grey or chestnut brown at centre, red-orange or orange near margin. Lamellae distant, L = 8–14, l = 0–2, collariate, white, whitish or cream, with pale brown or orange edge. Stipe 15–38 x 0.1 mm, filiform, glabrous, smooth, insititious, chestnut brown or chestnut brown-orange, then almost black. Context thin. Rhizomorphs often developed, black, glabrous, sometimes branched, thin. (According to Singer 1964, 1965a). — Pl. 1

 

Basidiospores 7.7–9.6(–10) x (3.8–)4.2–5.4 μm, E = 1.5–2.3, Q = 1.8–2.1, ellipsoid to broadly ellipsoid, thin-walled, hyaline, smooth. Basidia 20 x 6.9 μm, 4-spored, clavate. Basidioles 10–27 x 3.8-10 μm, clavate, cylindrical, subfusoid. Cheilocystidia in the form of broom-cells, 9.2–14.5 x 5.4–7.7 μm, clavate, thin-walled, with slightly thick-walled projections. Pleurocystidia not found. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, up to 8.0(–10) μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 11.5–15.5 x 6.2–12 μm, clavate to subcylindrical, entirely thin-walled or thin-walled below and slightly thick-walled above or entirely distinctly thick-walled, mixed with scattered, thick-walled, ± coralloid cells; projections 10–25, digitate, nodulose, obtuse, slightly thick-walled, up to 5.5 x 1.5 μm; thick-walled parts yellowish brownish in KOH. Pileocystidia absent. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6 μm wide hyphae, with yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 11

 

Chemical reactions. Stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing densely gregariously on dead leaves in a tropical forest.

 

Distribution. Collected only in Cameroon, Democratic Republic of Congo, Nigeria, Uganda and probably also Tanzania.

 

Revised specimens from tropical Africa.

Cameroon. ? Sud Province, Somalomo, Dja Biosphere Reserve, 8 Apr. 2001, V. Antonín Cm 01.46 (BRNM 666113).

Democratic Republic of Congo. Equateur Province, Eala, July 1907, L. Pynaert 1608 (BR 11515–69, holotype); Ibid., L. Pynaert 1607 (BR 11516–70); Ibid., 15 June 1907, L. Pynaert 1447 (BR 11517–71); Yambao, 19 June 1939, J. Louis 15235 (BR 11514–68); Kivu, Irangi, 19 April 1972, J. Rammeloo Z 335 (GENT); ? Tshopo Province, Kisangani, 8 April 1984, B. Buyck 1361 (BR 11767–30).

Nigeria. Cross River State, Anua Ranch, 14 April 1990, R.A. Nicholson 339 (K(M) 16767).

Tanzania. ? West Usambara Mts., Shagayu Forest Reserve, Mlalo Mission, 15 Febr. 1985, I. Krisai 3754 (WU); ? Tanga Province, Tanga District, Usambara Mts., Amani, 18–19 Feb. 1973, L. Ryvarden 10725 (K(M) 8879, as M. haematocephalus).

Uganda. ? Buganda Province, Mengo District, Mabira Forest, 9 June 1968, D.N. Pegler U 1353 (K(M) 115018).

 

Notes. Marasmius subruforotula is characterised by having a grey or chestnut brown pileus at centre, red-orange or orange near margin, distant lamellae (L = 8–14), often well-developed rhizomorphs, 7.7–9.6(–10) x (3.8–)4.2–5.4 μm basidiospores, small cheilocystidia (9.2–11.5 x 5.4–7.7 μm) and small pileipellis broom-cells (11.5–15.5 x 6.2–12 μm). In the original description by Singer (1964, 1965a), basidiospores are indicated as somewhat smaller (7–8.5 x 3.2–4.2 μm). Nevertheless, my microscopic studies fit well with those published by Pegler (1977). The collection V. Antonín Cm 01.46 from Cameroon is included with a question-mark here, because of slightly different basidiospores (7.0–9.0 x 3.5–4.7 μm) and concolorous central papilla at pileus centre; the collection B. Buyck 1361 differs by brown pileus (6D6–8 to 6E8) – other macroscopic and microscopic characters agree with other mentioned collections.

Also collection Pegler U 1353 from Uganda is included with a question-mark here. It differs by a larger, up to 10 mm broad, white striate pileus and larger cheilocystidia (9.0–20 x 6.0–10 μm), and also the basidiospores are slightly longer (9.5–11 x 4.5–5.5 μm). It may even represent a separate taxon.

This species belongs to a complicated group of very similar species: M. ruforotula Singer, M. acicularis Berk. & M.A. Curtis, M. rufomarginatus Singer, M. guyanensis Mont., M. praecox Singer and M. foliicola Singer. It differs from all of them in having a very distinct acute papilla. Moreover, M. rufomarginatus has narrower basidiospores (6.5–10 x 2.5–4.5 μm) (Singer 1976); M. foliicola has a longer stipe (15–75 mm long); M. praecox grows only on wood (Singer 1976); M. acicularis has smaller basidiospores (6–8 x 3.5–5 μm), two types of cheilocystidia and scattered caulocystidia (Desjardin & al. 2000); M. guyanensis has larger, lacrimoid or subfusoid basidiospores. In M. ruforotula Singer, Singer (1976) described both a concolorous and discolorous lamellar edge and narrow basidiospores 7.2–10.2 x 3.5–4.3 μm, Pegler (1983) a pale rusty brown lamellar edge and 8–11 x 4.5–6 μm basidiospores, Desjardin & Horak (1997) a concolorous edge and 7–9.5 x 4–5 μm basidiospores, and Desjardin & al. (2000) both a concolorous and pale brownish lamellar edge and (6–)7–10(10.5) x 3–5(–5.5) μm basidiospores. Moreover, the latter authors included also carpophores with greyish brown to brown pilei here.

 

 8. Marasmius conicopapillatus Henn.

Hennings, Bot. Jahrb. Syst. 22: 100 (1895). Type: Cameroon, Ekundu–Liongo, 20 May 1892, P. Dusén 41 (UPS, holotype ?). The type material consists of several broken carpophores (dried-up alcohol material).

 

Selected descriptions and icons. Dennis, Trans. Brit. Mycol. Soc. 34: 418 (1951); Desjardin et al., Sydowia 52: 124–126 (2000); Hennings, Bot. Jarhb. Syst. 22: 100 (1895); Pegler, Kew Bull. Addit. Ser. 6: 165 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 338–339 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 260 (1965).

 

Pileus 1–6 mm broad, convex, then plano-convex to applanate, umbilicate, with papilla in umbilicus, striate, white, then pale yellow to pale yellow-orange (± 4A3–5B3) with brown, then black-brown to black centre. Lamellae distant, L = 6–12, l = 0(–1), collariate, white, whitish or pale cream (4A2), with concolorous, finely pubescent edge. Stipe 10–40 x 0.1–0.2 mm, filiform, glabrous, smooth, insititious, lustrous, concolorous with lamellae above, straw coloured, soon darkening to dark brown or black-brown towards base. Context thin. Rhizomorphs and sterile stipes present. — Pl. 2

 

Basidiospores 7.3–10.5(–12) x 3.5–5.5 μm, E = 1.7–2.5, Q = 1.9–2.3, ellipsoid or sublacrimoid, thin-walled, smooth, hyaline. Basidia (16–)22–30 x 5.0–9.5 μm, 4-spored, clavate. Basidioles 12–30 x 3.5–9.0 μm, clavate, cylindrical or fusoid. Cheilocystidia similar to pileipellis cells, 10–20 x (5–)7.0–11(–13) μm, ± thin-walled, clavate, hyaline. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, branched up to 15 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, (7.0–)12–22 x 6.5–11.5 μm, clavate to subvesiculose, entirely thin-walled or slightly thick-walled above and in projections; projections digitate, obtuse to subacute, nodulose, slightly thick-walled, 1.5–8.0(–12) x 0.5–1.5 μm; mixed with entirely thick-walled broom-cells or coralloid cells; thick-walled parts pale yellowish brownish or greyish yellowish in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 5.0 μm wide hyphae with dark yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 12

 

Chemical reactions. Stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves in a tropical forest.

 

Distribution. A pantropical species. It has been described from Cameroon, and also collected in Burundi, Democratic Republic of Congo, Ghana, Ivory Coast, Nigeria, Uganda and Sierra Leone (Pegler 1977), Asia (Indonesia) (Desjardin & al. 2000) and South America (Bolivia, Trinidad, Jamaica) (Singer 1976).

 

Revised specimens from tropical Africa.

Burundi. ? Muramvya Province, Teza, 20 Dec. 1978, J. Rammeloo 6165 (BR 11909–75).

Democratic Republic of Congo. Eala, June 1923, M. Goossens–Fontana 109 (BR 11424–75); Yangambi, 22 May 1939, J. Louis 14935 (BR 11425–76); Kisantu, 20 March 1907, H. Vanderyst s.n. (BR 13866–92); Kivu, Irangi, 11 March 1972, J. Rammeloo Z 62 (GENT); without locality and date, H. Vanderyst 8270 (BR 13721–44); ? Tshopo Province, Kisangani, 8 April 1984, B. Buyck 1360 (BR 11768–31).

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 7 April 2001, V. Antonín Cm 01.39 (BRNM 666107); Ibid., 9 April 2001, V. Antonín Cm 01.67 (BRNM 666126); South–West Province, Korup National Park, transect P10, 22 March 1991, R. Watling s.n. (E); Ekundu–Liongo, 20 May 1892, P. Dusén 41 (UPS, holotype ?).

Ghana. Tafo, April 1955, M. Holden GC 38 (K(M) 11502, as M. graminum).

Ivory Coast. Forêt de la Besso, 31 March 1975, L. Aké Assi 322 (K(M) 111227, as M. baumannii).

Nigeria. Cross River State, Calabar-Cameroon Road, 3 July 1990, R.A. Nicholson 601 (K(M) 16697); Ibid., 23 June 1990, R.A. Nicholson 456 (K(M) 16571); Ibid., 4 Aug. 1990, R.A. Nicholson 694 (K(M) 16577); Cross River State, Uyo, Anua Ranch, 10 June 1989, R.A. Nicholson 225 (K(M) 7642); Akwa Ibom State, Anua Ravine, 13 May 1989, R.A. Nicholson 204 (K(M) 7502, as M. haediniformis).

Uganda. ? Buganda Province, Mengo District, N of Entebbe, Zika Forest, 12 June 1968, D.N. Pegler U 1431 (K(M) 111228, as M. baumannii); Buganda Province, Mengo District, Mawakota County, Mpanga Research Forest, 8 June 1968, D.N. Pegler U 1307 (K); Western Province, Bunyoro District, near Masindi, Budongo Forest, 15 June 1968, D.N. Pegler U 1472 (K).

Sierra Leone. Njala, Aug. 1949, F.C. Deighton M 3056 (K).

 

Notes. Marasmius conicopapillatus is characterised by having a white, then pale yellow to pale orange pileus with a brown to black papilla, distant lamellae (L = 6–12), a straw coloured stipe, soon darkening to dark brown or black-brown, small, 10–20 x (5–)7.0–11(–13) μm cheilocystidia and small, (7.0–)12–22 x 6.5–11.5 μm pileipellis cells. A description by Pegler (1977) differs by smaller basidia (13–21 x 5–6 μm), cheilocystidia (8–13 x 5–10 μm) and pileipellis cells (8.5–12 x 5–8 μm) and smaller basidiospores (7.5–9.5 x 3.3–4.3 μm). On the contrary, our description fully agrees with the one by Desjardin & al. (2000) except for the larger cheilocystidia (8–30.5 x 6–12 mm). The collections included with a question-mark are either without a description or with very sparse material but probably belong to this species. The type specimen differs in slightly smaller basidiospores (6.5–9.0 x 4.0–5.5(–6.0) µm).

Marasmius pallenticeps Singer, known from Argentina and New Zealand, has smaller basidiospores (8–10 x 4–5 μm) and its stipes arise directly from rhizomorphs (Desjardin & Horak 1997, Singer 1976); M. peckii Murrill, from Belize, differs by a smaller pileus (up to 2 mm broad), smaller basidiospores (5.8–6.8 x 1.8–2.8 μm) and smaller pileipellis broom-cells (5.5–10 x 5.5–7.8 μm); M. chrysochaetes (Berk. & M.A. Curtis) Berk., from Cuba, differs also in having a smaller pileus (1.2–2 mm broad) and basidiospores (8–9 x 3.5–4.2 μm), and M. aspilocephalus Singer, from Bolivia, in having a pale buff or pale cinnamon coloured pileus papilla, smaller pileipellis broom-cells (9–11 x 8–11 μm) and by growing on monocotyledons (bamboo) (Singer 1976).

 

9. Marasmius crinisequi F. Muell.

F. Mueller in Kalchbr., Grevillea 8: 153 (1880). Marasmius crinisequi var. monocotyledonum Singer, Fl. Neotropica 17: 148 (1976). Type: Australia, Queensland, Rockingham Bay, F. von Mueller s.n. (K(M) 99658, lectotype). – Marasmius equicrinis F. Muell. in Berk., J. Linn. Soc. Bot. 383 (1881).  – Androsaceus equicrinis (F. Muell.) Overeem, Hoofd Mus. Econ. Bot. Buitenzorg 1: 69 (1927). Marasmius graminum (Lib.) Berk. var. equicrinis (F. Muell.) Dennis, Trans. Brit. Mycol. Soc. 34: 416 (1951).  Marasmius repens Henn., Bot. Jahrb. Syst. 23: 548 (1897).

 

Selected descriptions and icons. Beeli, Bull. Jard. Bot. Etat Brux. 15: 37 (1938); Dennis, Trans. Brit. Mycol. Soc. 34: 416 (1951) (as M. graminum var. equicrinis); Desjardin & al., Sydowia 52: 126–128 (2000); Nicholson, Nigerian Field 54: 25 (1989); Pegler, Kew Bull. Addit. Ser. 6: 164–165 (1977); Pegler, Kew. Bull. Addit. Ser. 9: 203–204 (1983); Pegler, Kew Bull. Addit. Ser. 12: 151 (1986); Petch, Tr. Brit. Mycol. Soc. 31: 33–34 (1948) (as M. equicrinis); Singer, Bull. Jard. Bot. Etat Brux. 34: 337–338 (1964); Singer, Flore Icon. Champ. Congo 14: 260 (1965); Singer, Fl. Neotropica Monogr. 17: 148–149 (1976).

 

Pileus 1.5–3 mm broad, convex, then applanate, umbilicate, striate, glabrous, light brown to brown, then mostly pale orange-brown or reddish brown, strongly and distinctly papillate, papilla less distinct when old, very dark. Lamellae distant, L = 6–8, collariate, moderately large, sometimes not reaching the pileus margin, pale cream or yellow orange. Stipe 3–10 x 0.1–0.15 mm, filiform, glabrous, smooth, attached to rhizomorphs, chestnut-brown, then black. Context very thin. Rhizomorphs abundant, long, black, glabrous. (According to Pegler 1977 and Singer 1964, 1965a).

 

Basidiospores 7.7–12 x 4.2–6.2 μm, E = 1.5–2.4, Q = 1.8–2.1, ellipsoid, thin-walled, smooth, hyaline. Basidia 20–30 x 6.5–9.2 μm, 4-spored, clavate. Basidioles 11.5–29 x 2.7–9.0 μm, cylindrical, clavate, subfusoid. Cheilocystidia similar to pileipellis cells, 15.5–21(–26) x 5.4–11 μm, clavate or subcylindrical, thin-walled. Pleurocystidia absent. Pileipellis a hymeniderm of broom-cells of the Siccus-type, 12.5–23 x 7.7–12.5 μm, clavate or subcylindrical, thin-walled with slightly thick-walled upper part and projections, rarely entirely slightly thick-walled and then often coralloid; projections obtuse to subacute, nodulose, slightly thick-walled, yellow-brown in KOH, 1.2–6.2(–7.5) x 0.7–2.0 μm. Stipitipellis a cutis composed of cylindrical, parallel, slightly thick-walled, up to 5.0 μm wide hyphae, with dark yellow-brown walls in KOH. Caulocystidia absent; adpressed to erect terminal cells present. Clamp-connections present in all tissues. — Fig. 13

 

Chemical reactions. All structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead twigs and leaves of both dicotyledons (e.g. cacao-tree) and monocotyledons (e.g. bamboo). It causes a “Horse-hair blight” disease.

 

Distribution. Pantropical species; in Africa, it has been collected in Burundi, Cameroon, Democratic Republic of Congo, Ghana, Ivory Coast, Kenya, Nigeria and Sierra Leone. Moreover it is known from Australia (type), Asia (Indonesia, Malaysia, Sri Lanka) and tropical America (Guadaloupe, Martinique, Trinidad).

 

Revised specimens from tropical Africa.

Burundi. Muramvya Province, Bugarama, 19 Nov. 1966, J. Lewalle 1262 (BR 11432–83); Ibid., 14 Dec. 1967, E. Petit 2002 (BR 13735–58); Bururi Province, Bururi, 7 Feb. 1979, J. Rammeloo 6592 (BR 11932–01).

Cameroon. Bipinde, G. Zenker 129 (isotype of M. repens, S F–16262).

Democratic Republic of Congo. Angodia, Lebrun 3009 (BR 11433–84); Bamania, Aug. 1930, P. Staner 325 (BR 11434–85); Kivu, Irangi, 11 March 1972, J. Rammeloo Z 64 (GENT); ? Ibid., 11 March 1972, J. Rammeloo Z 56 (GENT); Kivu, Kahuzi volcano, 17 March 1972, J. Rammeloo Z 121 (GENT).

Ghana. Kibbi, 1922, W. Bunting s.n. (K(M) 135140).

Ivory Coast. Forêt de la Besso, 29 Nov. 1973, L. Aké Assi 167 (K(M) 135141).

Kenya. Nyanza province, Kericho District, Kericho, Kiptiget River, African Highland Estate, Man Forest, 26 March 1968, D.N. Pegler K 259 (K(M) 133704).

Nigeria. Ibadan, Maar Plantation, Nov. 1966, I.G. Weststeijn s.n. (K(M) 133701); ? Cross River State, Obudu Ranch, 25 April 1989, R.A. Nicholson 186 (K(M) 7471; ? Ibid., 25 April 1989, R.A. Nicholson 184 (K(M) 7506, as M. cupressiformis); ? Cross River State, Calabar-Cameroon Road, 23 June 1990, R.A. Nicholson 545 (K(M) 18619, as M. cupressiformis).

Sierra Leone. Njala, 20 Nov. 1935, F.C. Deighton M 888 (K(M) 133702); Ibid., 20 Nov. 1935, F.C. Deighton M 891 (K(M) 133703).

 

Revised specimens from other regions.

Australia. Queensland, Rockingham Bay, F. von Mueller s.n. (K(M) 99658, lectotype).

 

Notes. Marasmius crinisequi is characterised by a small, brown, later usually orange to reddish brown pileus with a distinct dark papilla, distant lamellae with a concolorous edge and a stipe inserted directly on rhizomorphs. Singer (1964, 1965a) described narrower basidiospores (7.5–10 x 3.7–5 mm), Pegler (1977, resp. 1983) narrower basidiospores (9–13 x 3.5–5.0 μm, resp. 9–11.5 x 3.5–4.5 μm), larger and smaller basidia (14–16 x 4.5–5.5 μm), Desjardin & al. (2000) narrower basidiospores ((7–)8–11(–12) x 3–5 μm). Collection J. Rammeloo Z 56 macroscopically differs by the absence of a coloured central papilla (there are no microscopical differences), however, a complete macroscopic description of fresh carpophores is not available. Therefore, it is included with a question mark here.

Singer (1976) described the new variety monocotyledonum Singer based on the growth on monocotyledons and basidiospores up to 11 μm long; collection Stanner 325 from the Democratic Republic of Congo was proposed as the type collection. However, both ecology and basidiospore size fall within the variability of typical M. crinisequi.

Marasmius pallenticeps Singer, known from Argentina and New Zealand, has a white pileus, more numerous lamellae (L = 9–10) and smaller basidiospores (8–10 x 4–5 μm) (Desjardin & Horak 1997; Singer 1976); M. berambutanus Desjardin, Retnowati & E. Horak, from Indonesia, has a longer stipe (4–25 mm long), slightly narrower lamellae ((7–)8–11 x 3–5 μm) and present pileosetae (Desjardin & al. 2000). Marasmius schultesii Singer, from Colombia, and M. hippiochaetes Berk., known from Surinam, Venezuela, Brazil and Bolivia, have hirsute-pilose rhizomorphs (Singer 1976); M. polycladus Mont., from French Guyana, has a brownish purple or blood red pileus and smaller basidiospores (6.3–7.7 x 2–3 μm); M. microdendron Singer, from Brazil, has a very small pileus (0.5–1 mm), which is pink or purplish pink, smaller basidiospores (7.5–8 x 2.8–3 μm) and sparsely hirsute rhizomorphs; M. robertsonii Singer, from Bolivia, has a larger, orange-ferrugineous coloured pileus, a longer stipe (7–20 mm) and smaller basidiospores (10 x 3.5–4.1 μm) (Singer 1976).

 

10. Marasmius curreyi Berk. & Broome var. distantifolius Antonín

Antonín, Mycotaxon 89(2): 425 (2004). Type: Benin, Oueme Province, Agongo, 17 Aug. 1997, V. Antonín B97.40 (BR 101094–20, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 89(2): 425-427 (2004).

 

Pileus 1.5–8 mm broad, hemispherical to convex with central umbilicus with a (distinctly projecting) papilla when young, then campanulate-convex to broadly convex, rarely almost applanate with papilla when old, sulcate, crenulate at margin, finely pubescent, slightly cracking around umbilicus when old, cinnamomeous brown to reddish (up to 8C7–8) when young, pallescent with age up to pinkish ochraceous (6A4–7A4), sometimes washed-up up to whitish at margin when old, always dark brown at centre. Lamellae distant, L = (6–)7–9, l = 0(–1), broad, sometimes almost ventricose when old, slightly intervenose at base when old, collariate, collarium often funnel-shaped when old, pale cream when young (paler than 5A2), sometimes with an ochraceous tinge when old, with (irregularly) red-brown, finely pubescent edge. Stipe 4–12 x 0.1–0.2 mm, filiform, lustrous, insititious, smooth and glabrous, concolorous with lamellae at apex, through brown to black-brown towards base. — Pl. 2

 

Basidiospores (8.0–)9.0–10.5(–12) x (4.5–)5.0–6.0 μm, E = 1.6–2.3, Q = 1.8, (broadly) ellipsoid, ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidia 24–28 x 8.0–10 μm, 4-spored, clavate. Basidioles 18–30 x 4.0–11 μm, cylindrical, clavate, fusoid. Cheilocystidia shaped as broom-cells of the Siccus-type, 10–23 x 6.5–12 μm, clavate, subcylindrical, thin-walled with slightly thick-walled, nodulose, obtuse, up to 10 x 2.0 μm projections; projections with slightly greyish yellowish walls in KOH. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, branched, thin-walled, smooth to minutely incrusted, up to 12 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, (5.0–)12–18 x 5.0–11(–13) μm, (broadly) clavate, subcylindrical, ± thin-walled, with slightly thick-walled, nodulose, digitate, obtuse projections, mixed with entirely or at least in upper part thick-walled, ± coralloid cells or broom-cells; thick-walled parts greyish ochraceous in KOH. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 6.0 μm wide hyphae with brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 14

 

Chemical reactions. Stipitipellis dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on grass remnants in open grassy place.

 

Distribution. Cosmopolitic species known from Europe, North and South Americas, Asia and Oceania; only one collection is known from tropical Africa (Pegler 1968).

 

Revised specimens from tropical Africa.

Benin. Oueme Province, Agongo, 17 Aug. 1997, V. Antonín B97.40 (BR 101094–20, holotype).

 

Notes. Marasmius curreyi var. distantifolius is especially characterised by coloured lamellar edge and distant lamellae. The closest taxon, var. culmisedus Singer, differs by slightly longer and narrower basidiospores ((8-)9-12.3 x (4-)4.2–5.3(-7) μm) and closer lamellae (L = 9–18) (Singer 1976, as M. graminum var. culmisedus Singer). Other varieties of the latter species differ by white coloured lamellar edge, differently numerous lamellae and/or different size of basidiospores (Singer 1976). For relation between M. curreyi and M. graminum, see Antonín & Noordeloos (1993).

Collection E. Harris 443 (Nigeria, Zaria Province, Shika, on leaf sheaths of Sorghum vulgare, 8 Sept. 1958, det. G.F. Laundon, as M. pruinatus, K(M) 110681) also is very close to M. curreyi by having ellipsoid, 9.0–12 x 5.0–6.0 μm basidiospores and moderately close lamellae (L = 9–12); the lamella edge seems to be concolorous and the carpophores darker (± brown-red or brown ferrugineous); a macroscopic description is not available.

Eichelbaum (1907) published a similar taxon (as M. graminum) from Tanzania (East Usambara). However, this fungus probably represents a different species.

 

11. Marasmius yangambiensis Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 335 (1964). Type: Democratic Republic of Congo, Yangambi, Ile Esali, 16 Dec. 1937, J. Louis 7091 (BR 11533–87, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 335–336 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 259 (1965).

 

Pileus 1.5–2.5 mm broad, convex, umbilicate, with low papilla in umbilicus, striate, glabrous, cinnamomeous brown (13–1–9/10, Maerz & Paul when dry), chestnut brown in papilla with a paler zone around. Lamellae rather distant, L = 10–12, collariate, rather large, white, with concolorous edge. Stipe 16–30 x 0.1 mm, filiform, glabrous, smooth, insititious, brown. Context thin. Rhizomorphs present, brown to brown-black. (According to Singer 1964, 1965a).

 

Basidiospores 16–26 x 4.2–5.8 μm, E = 3.2–5.1, Q = 4.2, clavate to lacrimoid, thin-walled, smooth, hyaline. Basidia not found. Basidioles 11.5–27 x 4.0–9.0 μm, cylindrical, clavate, subfusoid; some 24.5–33 x 8.5–13 μm, clavate, fusoid, thin-walled, hyaline “pleurocystioid basidioloid cells” present in hymenium. Cheilocystidia similar to pileipellis cells, 10–14.5 x 6.9–10 μm, clavate, thin-walled. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 7.7–14 x 5.4–10 μm, (broadly) clavate, rarely subcylindrical, entirely or only at apex slightly thick-walled, rarely entirely thin-walled, with thick-walled, slightly nodulose, obtuse to subacute, up to 4.0 x 1.2(–1.5) μm projections; thick-walled parts yellow-brown in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 6.5 μm wide hyphae, with yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 15

 

Chemical reactions. Stipitipellis hyphae slightly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing gregariously on dead twigs in a tropical forest.

 

Distribution. So far known only from the type locality in the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Yangambi, Ile Esali, 16 Dec. 1937, J. Louis 7091 (BR 11533–87, holotype).

 

Notes. Marasmius yangambiensis is especially characterised by having a cinnamomeous brown pileus, a brown stipe and very large basidiospores. The presence of such cells and the size of basidiospores more or less agree with sect. Sicci. However, the stipe is distinctly insititious and the lamellae are (also in exsiccates) collariate. Therefore, this species is placed in sect. Marasmius. Singer (1964, 1965a) mentioned smaller basidiospores (13.2–15.5 x 3.0–4.2 μm) and larger pileipellis cells (6.5–22 x 6.7–14 μm).

Among species from subsect. Sicciformis with large basidiospores, M. brevicolus Corner has a 3–9 mm broad, fuscous or subochraceus pileus and smaller (17–20 x 4.5–5.5 μm) basidiospores; M. adhaesus Corner has an up to 40 mm broad, greyish, fuligineous or brownish olive coloured pileus, a 30–45 x 1–2 mm stipe and larger (20–35 x 3–15 μm) cheilocystidia (Corner 1996); M. sanguirotalis Singer, from Argentina, has a purple coloured pileus with a purplish black centre, and smaller basidiospores (14.3–20 x 3–4 μm), and M. megalospermus Singer, from Bolivia, a brown to terracotta coloured pileus, a yellow succineous to pale sordid umbra coloured stipe and 16.5–19.5 x 4–4.2 μm basidiospores (Singer 1976) 

 

12. Marasmius aurantiostipitatus Antonín & P. Roberts

Antonín & P. Roberts in Antonín, Mycotaxon 89(2): 423 (2004). Type: Cameroon, South West Province, Korup National Park, trail from Rengo Camp to Erat, 2 May 1996, P.J. Roberts K 356 (K(M) 39176, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 89(2): 423-425 (2004).

 

Pileus up to 10 mm broad, campanulate, umbilicate, with a very small central papilla or without it, sulcate, smooth, brownish pink to dark pinkish red (maroon). Lamellae distant, L = 10–12, l = 0(–1), less distinctly collariate, broad, white, with concolorous edge. Stipe up to 45 x 0.5 mm, filiform, smooth, glabrous, lustrous, orange to deep orange. Sterile stipes and rhizomorphs present.

 

Basidiospores 13.5–18 x 4.5–7.5 μm, E = 2.4–3.5, Q = 3.0, lacrimoid to fusoid, thin-walled, hyaline, smooth. Basidia 30–33 x 7.5–10 μm, 4-spored, clavate. Basidioles 18–38 x 4.0–8.0 μm, clavate, subcylindrical, fusoid. Cheilocystidia in the form of broom-cells of the Siccus-type, 7.5–21 x 5.5–11 μm, clavate, subcylindrical, thin-walled, with thin- to slightly thick-walled, obtuse, up to 7.0 x 1.5 μm projections. Pleurocystidia absent. Trama hyphae cylindrical or subinflated, smooth, hyaline, up to 15 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–20 x 4.5–11 μm, clavate, pyriform or subcylindrical, slightly to distinctly thick-walled (up to 1.5 μm), with conical or cylindrical, thick-walled, obtuse, up to 8.0 x 1.5 μm projections; mixed with thick-walled broom-cells or coralloid cells; thick-walled parts with pale ochraceous-brown walls in KOH. Stipitipellis a cutis of consisting cylindrical, parallel, slightly thick-walled, smooth, up to 5.0 μm wide hyphae with dark brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 16

 

Chemical reactions. Stipitipellis hyphae slightly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing gregariously on dead leaves and twigs.

 

Distribution. So far known only from Cameroon.

 

Revised specimens from tropical Africa.

Cameroon. South West Province, Korup National Park, trail from Rengo Camp to Erat, 2 May 1996, P.J. Roberts K 356 (K(M) 39176, holotype); Ibid., trail from Rengo Camp to Ekunde-Kunde, 4 May 1996, P.J. Roberts K 494 (K(M) 39177).

 

Notes. Marasmius aurantiostipitatus is characterised by having a brownish pink to dark pinkish red pileus, distant, less distinctly collariate lamellae with concolorous edge, an orange to deep orange stipe, rather large basidiospores, cheilocystidia with obtuse projections and a pileipellis of slightly to distinctly thick-walled broom-cells with obtuse projections, mixed with thick-walled broom-cells or coralloid cells.

Marasmius guyanensis Mont. has a yellow orange, orange, brownish red or ferrugineous pileus, less numerous lamellae (L = 6–10), a black stipe and smaller basidiospores ((9.0–)10–13.5(–14) x (3.0–)3.5–4.5(–5.0) μm); M. lovedalensis Antonín & Verbeken has a smaller, 2–5 mm broad, orange pileus, black coloured stipe and smaller basidiospores 11–13(–14.5) x 4.5–6.0 μm. Marasmius marthae Singer, described from Argentina, with a similarly coloured pileus, has more numerous lamellae (L = 16–17) with a deep purple edge and narrower basidiospores ((8.3–) 14.5–16 x 4(–4.3) μm); M. xerampelinus Singer, described from Mexico, has a purple red or purple pileus, a shorter (10–19 x 0.3–0.4 mm) black stipe and smaller, (6–)8.5–11 x (4–)5.5–7.5 μm, basidiospores (Singer 1976). 

 

13. Marasmius guyanensis Mont.

Montagne, Ann. Sci. Nat. Bot., sér. 4, 1: 114 (1854). Type: French Guyana, Leprieur duplicate of holotype (K, ex herb. Berkeley, Desjardin & al. 2000); not studied. — Marasmius ochraceo-niger Henn., Bot. Jahrb. Syst. 30: 47 (1901).

 

Selected descriptions and icons. Dennis, Tr. Brit. Mycol. Soc. 34: 417 (1951); Desjardin & al., Sydowia 52: 121–122 (2000); Singer, Fl. Neotropica Monogr. 17: 146–147 (1976).

 

Pileus 1.5–7 mm broad, subhemisphaerical or convex, with central umbilicus, with small central papilla, dot or without it, distinctly striate-plicate, finely tomentose, crenulate at margin, light yellow orange (5A4–6), then orange, brownish red or ferrugineous (6A6, 6B7–8, 6D8, 7–8C7–8, 8C6), darker at centre, with concolorous to black-brown or brown central papilla. Lamellae distant, L = 6–10, l = 0(–1), collariate, broad, white when young, soon yellowish white (4–5A2–3), with concolorous, almost smooth edge. Stipe 10–35 x 0.1–0.2 mm, filiform, smooth, glabrous, lustrous, ± concolorous at apex, black-brown to black towards base. Sterile stipes and rhizomorphs present. — Pl. 2

 

Basidiospores (9.0–)10–13.5(–14) x (3.0–)3.5–4.5(–5.0) μm, E = 2.3–3.5, Q = 2.5–2.8, lacrimoid, subfusoid, thin-walled, hyaline, smooth. Basidia 20–26 x 8.5–10 μm, 4-spored, clavate. Basidioles 10–27 x 4.0–10 μm, clavate, ellipsoid, fusoid. Cheilocystidia shaped as broom-cells of the Siccus-type, 9.0–18 x 5.0–13 μm, (broadly) clavate, thin-walled, with slightly thick-walled, obtuse or subacute, nodulose, up to 5.0 x 1.0(–1.5) μm projections. Pleurocystidia absent. Trama hyphae cylindrical or subinflated, branched, smooth or minutely incrusted, up to 15 μm wide. Pileipellis a hymeniderm of broom-cells of the Siccus-type, 10–23 x 6.0–10 μm, (broadly) clavate, thin-walled, with slightly thick-walled upper part, sometimes entirely slightly thick-walled, with conical or subcylindrical, slightly thick-walled, obtuse to subacute, up to 3.0 x 1.0 μm projections; mixed with thick-walled broom-cells or coralloid cells; thick-walled parts with pale ochraceous-yellow walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 6.0 μm wide hyphae with dark brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 17

 

Chemical reactions. Stipitipellis hyphae slightly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing gregariously on dead leaves.

 

Distribution. Possibly a pantropical species. Known from Africa (Benin, Cameroon, Democratic Republic of Congo, Nigeria and Sierra Leone, probably widely distributed there), South America (French Guyana, Trinidad, Bolivia, Brazil, Venezuela), as well as Asia (Indonesia).

 

Revised specimens from tropical Africa.

Benin. Atlantique Province, Pahou, 15 Aug. 1997, V. Antonín B97.33 (BR 101088–14); Atacora Province, Tanugu, 8 Sept. 1997, V. Antonín B97.174 (BR 101212–41).

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 7 April 2001, V. Antonín Cm 01.38 (BRNM 666106); Ibid., 8 April 2001, V. Antonín Cm 01.47 (BRNM 666114); Ibid., 11 April 2001, V. Antonín Cm 01.81 (BRNM 666075); South–West Province, Korup National Park, Mundemba, transect P3, 24 March 1991, R. Watling s.n. (E); Ibid., transect P–P, Aug. 1990, R. Watling s.n. (E); South–West Province, Korup National Park, 7 Aug 1990, R. Watling s.n. (E); ? South–West Province, Korup National Park, trail to Mana Bridge, 15 Apr. 1997, P.J. Roberts K 1204 (K(M) 91514, BRNM 677271); ? South–West Province, Korup National Park, trail from Rengo Camp to Ekunde-Kunde, 4 May 1996, P.J. Roberts K 510 (K(M) 39178, as M. crinisequi); Bipinde, March 1899, G. Zenker 2014 (S F–16272, isotype of M. ochraceo-niger).

Democratic Republic of Congo. Tshopo Province, Kisangani, 8 April 1984, B. Buyck 1350 (BR 11770–33); Ibid., 8 April 1984, B. Buyck 1351 (BR 11769–32).

Nigeria. Ibadan, Nigerian College AST, June 1957, D.J. Hankler 5 (K(M) 133697); Akwa Ibom State, Anua Ravine, 27 May 1989, R.A. Nicholson 213 (K(M) 111501, as M. beelianus); Cross River State, Uyo, Anua, St. Luke´s Hospital, 27 June 1990, R.A. Nicholson 478 (K(M) 16846, as M. subruforotula); Cross River State, Calabar-Ibom Road, 17 July 1990, R.A. Nicholson 636 (K(M) 16703, as M. beelianus); Cross River State, Uyo-Calabar Road, 22 July 1990, R.A. Nicholson 651 (K(M) 16729, as M. beelianus); ? Cross River State, Anua, 19 June 1985, R.A. Nicholson 77 (K(M) 111275, as M. crinisequi); ? Ibadan, Ife Biological Gardens, 1967, M.H. Zoberi 374 (K(M) 111251, as M. graminum).

Sierra Leone. Mano (Dase), 7 Febr. 1954, F.C. Deighton M 5795 (K(M) 133696).

 

Notes. Marasmius guyanensis is characterised by having a light yellow-orange to orange, brownish red or ferrugineous pileus with a small central papilla, dot or without a papilla, distant lamellae (L = 6–10), large basidiospores ((9.0–)10–13.5(–14) x (3.0–)3.5–4.5(–5.0)) μm and small cheilocystidia (9.0–18 x 5.0–13 μm).

Although the type specimen has not been studied, my descriptions of African collections fit well with descriptions by Dennis (1951), Singer (1976) and especially by Desjardin & al. (2000).

According to macroscopic features, this species belongs to a complicated group of very similar species of the M. ruforotula/subruforotula group (see also notes on M. subruforotula). However, it differs from all of them in having large and subfusoid to lacrimoid basidiospores.

The collection P.J. Roberts K 321 (Cameroon, Korup National Park, trail to Rengo Rock, 1 May 1996, K(M) 39175, as M. crinisequi) may belong to this group of species. However, it has a larger pileus (6–14 mm in dry carpophores), a paler, ± stramineous-brown (never black) stipe arising from both substrate and rhizomorphs, slightly larger basidiospores (11.5–14(–15) x 4.1–5.6 μm) and smaller cheilocystidia (8.5–13.5 x 4.4–7.5 μm) and pileipellis broom-cells (8.5–13.5 x 5.2–8.5 μm). However, neither a detailed macroscopic description nor photographs are available.

The type revision of M. ochraceo-niger Henn. showed that its microscopic characters agreee well with M. guyanensis. Although Hennings (1901) described its pileus as ochraceous, I consider it identical with M. guyanensis.

  

14. Marasmius lovedalensis Antonín & Verbeken

Antonín & Verbeken in Antonín, Mycotaxon 88: 60 (2003). Type: Zimbabwe, Mvuma, Lovedale Ranch, hill north of road in front of “incubator” – 1930A4, 2 Febr. 1999, A. Verbeken 99–136 (GENT, holotype; BRNM 677246, isotype).

 

Selected descriptions and icons. Antonín & Verbeken in Antonín, Mycotaxon 88: 60–61 (2003).

 

Pileus 2–5 mm broad, convex, with crenulate margin, umbilicate at centre, with small central black dot, without distinct papilla, plicate-striate, orange (similar to M. curreyi). Lamellae moderately distant, L = 8–12, l = 0, collariate, broad, dark cream, with concolorous edge. Stipe 12–15 mm long, filiform, insititious, smooth, glabrous, entirely black. Sterile stipes present. — Pl. 2

 

Basidiospores 11–13(–14.5) x 4.5–6.0 μm, E = 2.1–2.9, Q = 2.5, fusoid, thin-walled, hyaline, smooth. Basidia 25–32 x 8.0–9.0 μm, 4-spored, clavate. Basidioles 16–32 x 5.0–10 μm, clavate, cylindrical, subfusoid. Cheilocystidia shaped as broom-cells of the Siccus-type, 15–20 x 7.0–11 μm, clavate, subcylindrical, thin-walled, upper part slightly thick-walled, with slightly thick-walled, rarely thin-walled, nodulose, subacute to obtuse, up to 5.0(–6.0) x 1.0 μm projections; thick-walled parts subhyaline to pale yellow-ochraceous. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, ± thin-walled, branched, hyaline, up to 12 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–20 x 7.0–12 μm, (broadly) clavate, subcylindrical, thin-walled in lower part, slightly thick-walled in upper part; projections slightly thick-walled, obtuse or subacute, nodulose, up to 6.0 x 1.0(–1.5) μm; mixed with some thick-walled cells transient to a coralloid shape; thick-walled parts with ochraceous yellow walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, cylindrical, slightly thick-walled, up to 5.0 μm wide hyphae with dark brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 18

 

Chemical reactions. Stipe medulla and cortex hyphae weakly to distinctly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead twigs in a hill miombo forest with Brachystegia glaucescens.

 

Distribution. So far known only in Zimbabwe.

 

Revised specimens from tropical Africa.

Zimbabwe. Mvuma, Lovedale Ranch, hill north of road in front of “incubator” – 1930A4, 2 Febr. 1999, A. Verbeken 99–136 (GENT, holotype; BRNM 677246, isotype). 

 

Notes. Marasmius lovedalensis is characterised by having distant lamellae, an orange coloured pileus without a central papilla (only with a central black dot), rather large and broad basidiospores, large basidia and rather small cheilocystidia and pileipellis cells.

A very close species seems to be M. guyanensis Mont. However, it has less numerous lamellae (L = 6–10), narrower basidiospores, smaller basidia (20–26 x 8.5–10 μm) and it grows on dead leaves. Marasmius gordipes Sacc. & Paol., from Sri Lanka, has an ochraceous brown or reddish brown pileus, a very long stipe (60–130 mm), and shorter basidiospores (9.5–11 x 4.5–6 μm) (Pegler 1986; Petch 1948); M. ruforotula Singer, known from South America, differs especially in having a well-developed central papilla at the pileus centre, smaller (8–11 x 4.5–6 μm, resp. 7–9.5 x 4–5 μm), ellipsoid basidiospores and smaller basidia (20–24 x 6–7 μm) (Pegler 1983, Desjardin & Horak 1997); M. dicotyledoneus (Singer) Singer, from Argentina, differs by a paler pileus colour, a shorter stipe (4–6 mm) and narrower basidiospores (9.5–12.3 x 3.5–4.7 μm) (Singer 1976).

 

15. Marasmius nigrobrunneus (Pat.) Sacc.

Saccardo, Syl. Fung. 11: 37 (1895). Androsaceus nigrobrunneus Pat., J. Bot. 5: 308. (1891). Type: Vietnam (Tonkin), Hanoi, Keso, 31 May 1890, Bon 4397 (FH, holotype). — Marasmius griseoviolaceus Petch, Tr. Brit. Mycol. Soc. 31: 42 (1948).

 

Selected descriptions and icons. Desjardin & Horak, Bibl. Mycol. 168:71–72 (1997); Nicholson, Nigerian Field 54: 26 (1989); Patouillard, J. Bot. 5: 308 (1891); Pegler, Kew Bull. 21: 527 (1968) (as M. griseoviolaceus); Pegler, Kew. Bull. Addit. Ser. 9: 206 (1983); Pegler, Kew Bull. Addit. Ser. 12: 150–151 (1986); Pegler & Calonge, Bol. Soc. Micol. Madrid 22: 52 (1997); Petch, Tr. Brit. Mycol. Soc. 31: 42 & Pl. IV, fig. 14 (1948); Singer, Fl. Neotropica Monogr. 17: 128–129 (1976).

 

Pileus 2–9 mm broad, hemispherical to convex, with a central papilla, glabrous, entirely sulcate to deeply sulcate, with a low black papilla or central dot in the umbilicus or shallow depression, grey (between “Deauville sand” and “turtledove”, “smoke brown”), becoming “limestone” M&P, sometimes whitish cinereous tinged. Lamellae distant, L = 8–13, l = 0–1(–2), collariate, moderately broad to broad, white, with concolorous or grey-brown edge. Stipe 13–73 x 0.1–0.5 mm, filiform, unpolished to somewhat shining, glabrous, smooth, insititious, black or deep umber with white apex; arising from rhizomorphs or directly from the substratum. Context white, thin, inodorous. (According to Singer 1976).

 

Basidiospores 8.0–11.5 x 5.0–6.0(–6.5) μm, E = 1.6–2.0, Q = 1.7–1.8, (broadly) ellipsoid, ellipsoid-fusoid, smooth, thin-walled, hyaline. Basidia not found. Basidioles 15–28 x 5.0–9.0 μm, clavate, cylindrical or fusoid. Cheilocystidia similar to the pileipellis cells, 9.0–19 x 4.5–7.0 μm, clavate, thin-walled with slightly thick-walled apex and projections; thick-walled parts sometimes with brownish walls in KOH. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, up to 12 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–22 x 7.0–12 μm, clavate to subcylindrical, thin-walled with slightly or distinctly thick-walled apex, or entirely thick-walled, or with subcoralloid or irregular thick-walled cells, with brown walls in KOH; projections slightly to distinctly thick-walled, obtuse. Pileocystidia absent. Stipitipellis a cutis consisting of parallel, cylindrical, thick-walled, up to 6.0 μm wide hyphae with brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. — Fig. 19

 

Chemical reactions. Stipe medulla and cortex hyphae and some trama hyphae weakly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves and twigs of bamboo (African collections), grasses and other monotyledons, found also on Bromeliaceae and palms.

 

Distribution. In Africa, so far known only from Nigeria and Sierra Leone. However, it represents a pantropical species, and has been collected in Argentina, Bolivia (Singer 1976), India (Singer 1976), Papua New Guinea (Desjardin & Horak 1997), Sri Lanka (Pegler 1986), Trinidad (Dennis 1951; Pegler 1983), Venezuela (Pegler & Calonge 1997) and Vietnam (Patouillard 1891; Pegler 1983).

 

Revised specimens from tropical Africa.

Nigeria. Cross River State, Uyo, 17 May 1985, R.A. Nicholson 44 (K(M) 115026); Cross River State, Uyo, Anua, St. Lukes Hospital, 7 May 1990, R.A. Nicholson 428 (K(M) 16690).

Sierra Leone. Kori, Njala, 29 July 1941, F.C. Deighton M 2935 (K(M) 115025, as M. griseoviolaceus).

 

Notes. Marasmius nigrobrunneus is characterised by having a dark coloured pileus, a dark coloured lamella edge, a very long stipe, rather large and broad basidiospores and very dark pigmented pileipellis cells.

The microscopic description agrees with those published by Desjardin & Horak (1997), Pegler (1983, 1986) and Singer (1976). Only Singer (1976) described the presence of small, dark coloured hairs, which I have not seen in African material. Fungi collected by Desjardin & Horak (1997) had no rhizomorphs.

Species with a similar pileus colour are M. fuligineorotula Singer, from Bolivia, with larger basidiospores ((7.5–)10.2–12.3 x 4.5–6 μm) and growing on dicotyledons, and M. magnisetulosus Singer, from Venezuela, which has smaller basidiospores (5.5–6.5 x 3–3.5 μm), shorter basidia (18 x 5.5 μm) and a different pileipellis structure.

 

Sect. Hygrometrici Kühner

 

Marasmius sect. Hygrometrici Kühner, Botaniste 25: 95 (1933) (as Hygrometriceae).

– Type species: Marasmius hygrometricus (Brigant.) Sacc. [= M. corbariensis (Roumeg.) Singer].

 

Basidiocarps small to very small. Pileus membranaceous, usually dark coloured, often radially striate or grooved. Lamellae vein-like to well developed; hymenophore sometimes smooth when young. Stipe insititious, thin, dark coloured, sometimes absent. Rhizomorphs and sterile stipes absent or present.

 

Basidiospores ellipsoid, fusoid-ellipsoid to lacrimoid. Cheilocystidia present, often in more types. Pleurocystidia present or absent. Pileipellis hymeniform composed of broom-cells of the Rotalis-type, sometimes some cells smooth (never all of them). Pileocystidia present or absent. Hyphae non-dextrinoid.

 

Key to tropical African species

 

1.   Pileus white, with yellowish centre when old ..........................................................  21. M. subalbidulus

1*. Pileus distinctly coloured ......................................................................................................................... 2

 

2.  Pileus strongly echinulate-spinulose; pileipellis with chains of thick-walled cells .......... 16. M. thwaitesii

2*. Pileus never strongly echinulate-spinulose; pileipellis without chains of thick-walled cells..................... 3

 

3. Pileocystidia absent .................................................................................................................................. 4

3*. Pileocystidia present ................................................................................................................................ 5

 

4.  Cheilocystidia of one type; stipitipellis hyphae not diverticulate .............................. 19. M. parviconicus

4*. Cheilocystidia of two types; stipitipellis hyphae diverticulate ................................... 20. M. mulanjensis

 

5. Basidiospores (11.6–)12.5–16.2 x 3.5–4.2 μm, E = 3.1–4.6, Q = 3.6; stipe up to 30 mm long .............................. 18. M. nyikae

5*. Basidiospores smaller (8.5–)9.0–12.5 x 3.7–6.0 μm, E = 1.6–2.9, Q = 2.0–2.5; stipe shorter, up to 13 mm long ......... 6

 

6. Basidiospores (9.0–)9.5–11(–11.5) x 3.7–4.5(–5.0) μm, Q = 2.5; basidia 18–23 x 6.5–8.5 μm ........... 17.2. M. minutoides var. angustisporus

6*. Basidiospores broader, (8,5–)9.5–12.5 x 4.5–6.0(–6.2) μm, Q = 2.0; basidia 22–27 x 9.0–10.0 μ.... 17.1. M. minutoides var. minutoides

 

Species descriptions

 

16. Marasmius thwaitesii Berk. & Broome

Berkeley & Broome, Journ. Linn. Soc., Bot. 14: 39 (1873). Type: Sri Lanka, Kandy District, Peradeniya, Oct. 1868, G.H.K. Thwaites 827 (K(M) 99662, holotype).   Marasmius echinosphaerus Singer, Bull. Jard. Bot. Etat Brux. 34: 325 (1964). 

 

Selected descriptions and icons. Pegler, Kew Bull. 23: 245–247 (1968) (as M. echinosphaerus); Pegler, Kew Bull. Addit. Ser. 12: 153–154 (1986); Petch, Tr. Brit. Mycol. Soc. 31: 32 (1948); Singer, Bull. Jard. Bot. Etat Brux. 34: 325–326 (1964) (as M. echinosphaerus); Singer, Flore Icon. Champ. Congo, fasc. 14: 255 (1965) (as M. echinosphaerus).

 

Pileus 1.5–3 mm broad, 1.5–5 mm high, (sub)hemispherical, often higher than broad, then convex, inflexed at fimbriate margin, strongly echinulate-spinulose (“spines” up to 1.5 mm long), dry, hygrophanous, dark reddish chestnut brown (Singer: 15–E–12, Maerz & Paul) when dry, sulcate at margin. Lamellae distant, L = 10–17, l = 0, (almost) free, broad, white to whitish, edge dark brown. Stipe 3–35 x 0.2–1 mm, cylindrical, filiform, insititious, finely echinulate-floccose under lens, especially at base, glabrescent; concolorous with pileus, whitish at apex when young, then almost uniformly chestnut brown. (According to Singer 1964, 1965a and Pegler 1966).

 

Basidiospores 6.5–10 x 3.3–4.5(–5.5) μm, E = 1.6–2.1, Q = 1.9, ellipsoid, subfusoid to amygdaliform, hyaline, thin-walled (according to type specimen of M. thwaitesii, type specimen of M. echinosphaerus is sterile). Basidia 19–26 x 5.0–8.5 μm, 4-spored, clavate. Basidioles 15–31 x 3.5–8.0 μm, clavate to subcylindrical. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, 18–26 x 7.0–11.5 μm, clavate, thin-walled at base, slightly thick-, rarely thin-walled above (up to 2.0 μm), with (sub)hyaline to brown walls in KOH, projections up to 2.0 μm long, (sub)acute; (2) lageniform to fusoid, often rostrate, obtuse cells, 24–44 x 6.0–8.5 μm, thin- to slightly thick-walled, smooth. Pleurocystidia present, similar to cheilocystidia of type 2. Trama hyphae cylindrical to subinflated, thin-walled, in subpileipellis slightly thick-walled, hyaline, 1.5–10 μm wide. Pileipellis a hymeniderm composed of more types of cells: (1) broom-cells of the Rotalis-type, 18–36 x 9.0–19 μm, clavate, sometimes slightly irregular, thick-walled (up to 7.5 μm), brown in KOH, projections up to 2.0 μm long, (sub)cylindrical, digitate, mostly obtuse, sometimes (sub)acute; (2) chains of 11.5–50 x 5.5–12 μm, fusoid, limoniform, ellipsoid, rarely subcylindrical cells, with mostly rostrate terminal cells, thick-walled (up to 1.5 μm), with similar projections like pileipellis cells. Pileocystidia 21–27 x 6.9–7.7 μm, lageniform to subfusoid, rostrate, mostly subcapitate, thick-walled, pale to dark brown in KOH. Circumcystidia present, 57–63 x 10–14 μm, (sub)clavate, thick-walled, with numerous small wart-like projections. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 4.5 μm wide hyphae, with brown incrusted walls in KOH. Caulocystidia absent; stipe surface covered by a brown amorphous layer and a layer of thick-walled hyphae of the same colour. Clamp-connections present in all tissues. – Fig. 20

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. Gregarious, on dead herbaceous stems (holotype specimen), on wood in a “muhulu” vegetation and on Gmelina arborea (collections by Schmitz-Levecq and Odeyinde).

 

Distribution. Known from the Democratic Republic of Congo and Nigeria. Outside tropical Africa, it was collected in Asia (Sri Lanka) (Pegler 1986, Petch 1948), South America (Argentina) (Singer 1976), and South Africa (Doidge 1950).

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Shaba Province, Kipopo, 2 April 1959, M.C. Schmitz–Levecq 169 (BR 11436–87, holotype of M. echinosphaerus).

Nigeria. Northern Nigeria, Ilorin, 2 Nov. 1966, M.A. Odeyinde 280 (K(M) 11240).

 

Revised specimens from other regions.

Sri Lanka. Kandy District, Peradeniya, Oct. 1868, G.H.K. Thwaites 827 (K(M) 99662, holotype).

 

Notes. Marasmius thwaitesii is characterised by having a distinctly echinulate-spinulose pileus, rather small basidiospores, two types of cheilocystidia, well-developed pleurocystidia and, particularly, chains of fusoid, limoniform, ellipsoid, rarely subcylindrical, diverticulate, thick-walled cells.

It belongs to the small group of species having developed chains of thick-walled cells. Marasmius magnoliae Singer differs by a pileus with a small papilla, pallescent at margin, fimbriate-ciliate only at margin when young (Singer 1976); M. kroumirensis (Pat.) Sacc. & P. Syd. has a uniformly coloured, broadly umbonate pileus, more distant lamellae, shorter basidia (16.5–19.5 x 5–6 μm), and only one type of cheilocystidia (Pegler 1966). Moreover, according to Desjardin & al. (1992), both of them have just a granulose to pruinose pileus (except for the sometimes hairy-fimbriate margin) and a different microscopical structure of the pileus ornamentation. A very close taxon was described as Marasmius sp. by Desjardin & al. (1992) from the Hawaiian Islands. It is similar in having a pileus with spinulose to pyramidal outgrowths with a similar microscopic structure and by the presence of circumcystidia. However, it has a smaller (0.7 mm broad), dark brown coloured pileus, a smaller stipe (2 x 0.25 mm), long-celled, setulose marginal hairs and no pleurocystidia.

This species is also known as M. echinosphaerus Singer. However, Pegler (1986) synonymised it with M. thwaitesii, which represents the oldest name for this taxon.

Reid (1975) identified M. echinosphaerus identical with M. actiniceps (Kalchbr. & Cooke) D.A. Reid [= Mycena actiniceps (Kalchbr. & Cooke) Sacc.; Agaricus actiniceps Kalchbr. & Cooke]. However, I studied the type specimen (South Africa, Cape Province, M.C.Cooke, K(M) 92579!) which represents quite a different species not belonging to the genus Marasmius having a non-hymenidermal pileipellis; it also differs e.g. by distinctly larger basidiospores (10–12.5 x 4.5–5.5 μm). 

 

17. Marasmius minutoides Antonín

Antonín, Mycotaxon 85: 121 (2003). 

 

17.1. M. minutoides var. minutoides

Type: Rwanda, road Butare-Cyangugu, km 94, Forêt de Rugege, 31 Aug. 1974, J. Rammeloo Z 433 (GENT, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 121–123 (2003).

 

Pileus 1–3 mm broad, convex, applanate to slightly depressed at centre, without central papilla, inflexed, then straight to slightly reflexed when old at crenulate and tomentose-pubescent margin, sulcate, translucently striate, slightly tomentose to granulose; entirely bronze to dark brown (5E6, 7E–F8) except for paler sulci when young, soon pallescent up to beige, greyish orange to orange-grey (5B–C4, 6B2) towards margin, sulci remaining paler, almost dirty whitish. Lamellae distant, L = 6–9, l = 0–1, shortly adnate, without collarium, slightly intervenose at base when old, sometimes furcate; yellowish white (c. 3A2), edge entire, concolorous, slightly pubescent. Stipe 3–12 x ca. 0.3 mm, thin-cylindrical, slightly broadened at apex, with a minute disc at base, insititious, slightly to distinctly curved, smooth, except for slightly pruinose apex; concolorous with lamellulae (yellowish white, 3A2) above, black-brown towards base. Forming sterile stipes and rhizomorphs— Pl. 3

 

Basidiospores (8.5–)9.5–12.5 x 4.5–6.0(–6.2) μm, E = 1.6–2.5, Q = 2.0, (sub)fusoid to ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidia 22–27 x 9.0–10.0 μm, 4-spored, (broadly) clavate. Basidioles 16–27(–31) x 4.0–12.0 μm, clavate, cylindrical, often fusoid. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, (8.0–)14–23 x (4.5–)7.0–13 μm, clavate, broadly clavate, (sub)vesiculose, thin- to slightly thick-walled (0.5 μm) at base, thick-walled above, ± hyaline, thick-walled parts with pale yellowish-greyish walls in KOH, projections up to 1.5(–2.0) μm long, pale yellowish-greyish; (2) lageniform to fusoid, often rostrate, obtuse to subcapitate, 23–33 x 5.0–9.0 μm, ± thin-walled, with hyaline to pale yellowish greyish walls in KOH. Pleurocystidia present, scattered, similar to cheilocystidia of type 2. Trama hyphae ± cylindrical, thin- to slightly thick-walled, branched or with lateral (sometimes up to subcoralloid) projections, smooth or minutely incrusted, hyaline, up to 9.0 μm wide. Pileipellis a hymeniderm composed of clavate, ellipsoid to vesiculose broom-cells of the Rotalis-type, 13–25 x 7.0–15(–20) μm, thin-walled at base, thin- to thick-walled (up to 3.0 μm) above, warts ± narrowly conical, subacute to acute, up to 1.5(–2.0) μm long; thin-walled cells subhyaline to pale brownish with yellow-brown to yellow-grey projections, thick-walled ones dark (yellow-)brown in KOH. Pileocystidia 17–25 x 5.0–9.0 μm, lageniform to fusoid, often rostrate, obtuse to subcapitate, thin- to mostly slightly thick-walled, smooth or with some projections in basal parts, subhyaline, with yellow-grey to yellow-brown walls in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, thick-walled (up to 1.5 μm), diverticulate, up to 5.0(–6.0) μm wide hyphae, with brown walls in KOH; diverticula up to 4.0 x 1.0 μm, cylindrical to conical, obtuse to subacute, thin- to slightly thick-walled, pale brown to dark (yellow-) brown in KOH. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 21

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid or amyloid.

 

Ecology. Growing on dead stem of a liana, dead leaves and especially petioles of Mumulopsis in a mountain rain forest.

 

Distribution. Known from Benin and Rwanda.

 

Revised specimens from tropical Africa.

Benin. Atacora Province, Tanugu, Chutes de Tanugu, 8 Sept. 1997, V. Antonín B97.171 (BR 101209–38).

Rwanda. Road Butare-Cyangugu, km 94, Forêt de Rugege, 31 Aug. 1974, J. Rammeloo Z 433 (GENT, holotype).

 

Notes. Marasmius minutoides is characterised by having a small sulcate pileus coloured dark brown at centre, yellow-brown at margin and in sulci, rather large spores, two types of cheilocystidia, well-developed pleuro- and pileocystidia and no caulocystidia.

A similar species, Marasmius minutus Peck, known from the Northern Hemisphere (Europe, North America, Far East), has a non-hygrophanous, not translucently striate, pale red coloured pileus with darker centre, almost free to narrowly adnexed, vein-like lamellae, smaller basidiospores [6.0–9.5 x (2.0–)2.7–4.0(–5.0) μm], a pileipellis composed of two types of cells and well-developed caulocystidia (Antonín & Noordeloos 1993). Marasmius corbariensis (Roumeg.) Singer has a larger, 1–7 mm broad, glabrous, only slightly grooved pileus, closer lamellae (L = 10–14), a longer stipe (8–30 mm), smaller basidiospores ((8.0–)8.5–10 x 3.5–5.0(–5.5) μm) and no pileocystidia; it is widely distributed in Mediterranean Europe and North Africa, it has once been reported from tropical Africa from the former Zaire (Democratic Republic of Congo) (Hendrickx 1948).

Marasmius parviconicus Pegler (Pegler 1982) from Zambia has a pale brown pileus (but darker at centre and on the sulci), only one type of cheilocystidia (fusoid) and no pileocystidia. According to the type revision (Pegler 1966), the North African (Tunesian) species M. kroumirensis (Pat.) Sacc. & Syd. has a uniformly fuscous, broadly umbonate pileus, a reddish brown stipe, smaller basidia (16.5–19.5 x 5–6 μm), only one type of cheilocystidia, well-developed pleurocystidia and no pileocystidia; according to Pegler the type material is in extremely poor condition and he has not found any spores. However, the same author (Pegler 1986) compared it with M. micraster Petch, a species possessing two types of cheilocystidia.

According to Desjardin & Horak (1997), M. fishii G. Stevenson & G.M. Taylor, known from New Zealand, differs especially in having a glabrous pileus not paler coloured in sulci, larger basidiospores (11.5–13 x 6.5–7.0 μm) and by the absence of pleuro- and pileocystidia; M. exustoides Desjardin & E. Horak, also from New Zealand, has a glabrous, only at margin plicate pileus, larger pileocystidia (30–50 x 10–20 μm) and no pleurocystidia; M. micraster Petch, known from Sri Lanka, Malaya and New Zealand, has a glabrous, rusty brown to sooty brown pileus becoming orange to pale orangish brown when dry (but yellowish brown to blackish brown and paler in sulci, Pegler 1986, Petch 1948), larger basidia (18–30 x 7–9 μm, but only 16–18 μm long according to Pegler 1986) and no pleuro- and pileocystidia.

Marasmius ilicis Singer, from South America, has an orange brown to chestnut brown pileus sometimes pale to saffron orange in sulci and at margin, and smaller basidiospores (6.7–9.5 x 3.5–5 μm); also M. exustus Berk. & M.A. Curtis has smaller basidiospores (5.5–9.2 x 3–4.5 μm, Singer 1976).

17.2. M. minutoides var. angustisporus Antonín

Antonín, Mycotaxon 85: 123 (2003). Type: Benin, Oueme Province, Agongo, 17 Aug. 1997, V. Antonín B97.44 (BR 101098–24, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 123–124 (2003).

 

Pileus 1–2 mm broad, almost campanulate-convex when young, then convex to applanate-convex when old, crenulate and sometimes slightly translucent at margin, distinctly sulcate, pubescent, hygrophanous; dark brown (7F4) when young, pallescent up to yellow-brown towards margin, remaining dark brown at centre, the yellowish-brownish colour remains also in sulci. Lamellae distant, L = 7–8, l = 0(–1), broadly adnate; yellowish white (c. 3A2), edge entire, concolorous, slightly pubescent. Stipe 11–13 x up to 0.5 mm, filiform, insititious, smooth, glabrous, slightly pruinose at apex, slightly lustrous, whitish to slightly brownish above, dark brown (7F7) towards base.

 

Basidiospores (9.0–)9.5–11(–11.5) x 3.7–4.5(–5.0) μm, E = 2.0–2.9, Q = 2.5, fusoid to ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidia 18–23 x 6.5–8.5 μm, 4-spored, clavate. Basidioles 11–25 x 4.0–9.0 μm, fusoid, clavate, often (sub)rostrate. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, 12–25 x 9.0–17 μm, clavate, broadly clavate, vesiculose, thin-walled at base, slightly thick-walled above (walls up to 1.5 μm), ± hyaline, projections up to 1.5(–2.0) μm long; thick-walled parts and projections pale yellowish-greyish; (2) lageniform to fusoid, often rostrate, obtuse to subcapitate, 26–32 x 6.5–10.0 μm, ± thin-walled, smooth cells, with hyaline to pale yellowish greyish walls in KOH. Pleurocystidia present, similar to cheilocystidia of type 2. Trama hyphae ± cylindrical, often irregular to branched, ± slightly thick-walled, smooth to incrusted, hyaline to pale yellowish, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 13–25 x 8.0–20 μm, clavate, ellipsoid, globose to vesiculose, ± thin-walled below, thin- to slightly thick-walled (1.0–1.5 μm) above, thin-walled parts ± hyaline, thick-walled parts with pale yellow-grey-brown walls in KOH; warts up to 1.5(–2.0) μm long, cylindrical to conical, subacute to acute, warts pale yellow-grey-brown in KOH. Pileocystidia 20–25 x 5.0–7.0 μm, lageniform to fusoid, often rostrate, obtuse to subcapitate, thin- to slightly thick-walled, smooth or with some projections in basal parts, hyaline to brownish in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, thick-walled, diverticulate, up to 5.0 μm wide hyphae, with dark yellow-brown to brown walls in KOH; diverticula up to 3.0 x 1.0 μm. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 21

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. On dead leaves of raffia in a marshy forest with dominating raffia.

 

Distribution. Known only from the type locality in Benin.

 

Revised specimens from tropical Africa.

Benin. Oueme Province, Agongo, 17 Aug. 1997, V. Antonín B97.44 (BR 101098–24, holotype).

 

Notes. Marasmius minutoides var. angustisporus differs from the type variety in having narrower basidiospores, and shorter and narrower basidia.

 

18. Marasmius nyikae Antonín

Antonín, Mycotaxon 85: 126 (2003). Type: Malawi, Nyika National Park, Gîte Chelinda, 4 Dec. 1981, J. Rammeloo 7646 (BR 11987–56, holotype). 

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 126–128 (2003).

 

Pileus up to 4 mm broad, convex-hemispherical to broadly conical (paraboloid), sulcate, finely tomentose, grey- to black-brown, sometimes (old carpophores) radially striped. Lamellae distant, L = ± 10, narrow, possibly with a dark edge. Stipe up to 30 mm long, filiform, insititious, finely pruinose when young, then glabrous and smooth, lustrous, paler at apex, black-brown towards base; forming sterile stipes and rhizomorphs. (According to slide and exsiccate.) — Pl. 3

 

Basidiospores (11.6–)12.5–16.2 x 3.5–4.2 μm, E = 3.1–4.6, Q = 3.6, fusoid to narrowly ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidia 24–32 x 6.9–9.2 μm, 4-spored, clavate. Basidioles 15–31 x 6.9–9.2 μm, clavate, (sub)cylindrical, subfusoid. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, 17–31 x 7.0–11.5 μm, clavate to broadly clavate, thin- to very slightly thick-walled above, hyaline at base, brownish above; projections variable in number, up to 1.0 μm long, subacute, less frequently obtuse, brown in KOH; (2) lageniform or fusoid, 27–35 x 6.0–9.0 μm, thin- to slightly thick-walled, rostrate, subacute, rarely acute, obtuse or subcapitate cells. Pleurocystidia similar to cheilocystidia of type 2, 24–36 x (5.5–)6.0–7.5 μm, lageniform, subulate or (sub)fusoid, sometimes subcapitate, thin-, rarely slightly thick-walled. Trama hyphae cylindrical, often branched (sometimes up to coralloid), hyaline, smooth, up to 8.0 μm wide. Pileipellis a hymeniderm composed of more types of cells: (1) broom-cells of the Rotalis-type, (13–)14.5–32 x 8.0–13.5(–15.5) μm, clavate, broadly clavate to (sub)vesiculose, thin- to thick-walled (especially at the top); thick-walled parts with brown walls in KOH; projections wart-like or digitate, obtuse to (sub)acute, up to 1.5 μm long, brown in KOH; (2) scattered thick-walled cells of similar size and colour as type 1, but (almost) smooth. Pileocystidia 20–29 x 4.5–7.0 μm, lageniform or (sub)fusoid, rostrate, often (sub)capitate, thin- to slightly thick-walled, mostly smooth, rarely with some digitate projections at basal part. Stipitipellis a cutis consisting of cylindrical, parallel, thick-walled (walls up to 1.0 μm), up to 5.0 μm wide hyphae, with brown walls in KOH; stipitipellis hyphae diverticulate at apex when young, smooth towards base; diverticula up to 4.0 x 1.0 μm, digitate, obtuse. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 22

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid or amyloid.

 

Ecology. Growing on dead leaves at an altitude of 2300 m.

 

Distribution. Known only from Malawi.

 

Revised specimens from tropical Africa.

Malawi. Nyika National Park, Gîte Chelinda, 4 Dec. 1981, J. Rammeloo 7646 (BR 11987–56, holotype).

 

Notes. Marasmius nyikae is characterised by having a dark coloured, sometimes striped pileus, large fusoid to ellipsoid-fusoid basidiospores, two types of cheilocystidia and present pleuro- and pileocystidia. Such a combination of microfeatures has not been found in any other species of this section.

 

19. Marasmius parviconicus Pegler

Pegler, Kew Bull. 37(2): 260 (1982). Type: Zambia, Kitwe, 5 Dec. 1978, G.D. Piearce FP 591 (K(M) 99655, holotype). 

 

Selected descriptions and icons. Pegler, Kew Bull. 37(2): 260–261 (1982).

 

Pileus 3–5 mm broad, conical to broadly campanulate with depressed disc, surface pale brown, much darker on disc and on the striae, plicate, pruinose. Lamellae adnexed to adnate, not collariate, white, narrow, distant, 8–14 entire, with occasional lamellulae; edge white. Stipe 7–12 x 0.3–0.5 mm, filamentous, cylindrical, solid; surface brown at apex, black elsewhere, glabrous, smooth and shiny, insititious. Context membranaceous, white. Odour none. (According to Pegler 1982). — Pl. 3

 

Basidiospores (only 5 found) 9.0–11 x 4.0–5.5 μm, ellipsoid-lacrimoid, subfusoid, thin-walled, smooth. Basidia 25 x 10.5 μm, 4-spored, clavate. Basidioles 15–28 x 6.0–11 μm, clavate to fusoid, then rostrate. Cheilocystidia 29–38 x 8.0–13 μm, fusoid, sometimes rostrate, thin-walled. Pleurocystidia similar to cheilocystidia. Trama hyphae cylindrical, subinflated, thin-walled, mostly smooth, rarely finely incrusted, hyaline, up to 8.0(–10) μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 14–30 x 8.0–14(–20) μm, clavate, broadly clavate, (sub)vesiculose, thin- or sometimes slightly thick-walled at base, slightly to distinctly thick-walled above (walls 0.5–2.0(–5.0) μm); diverticula wart-like to digitate, up to 2.0 μm long; thick-walled parts and projections rarely with subhyaline, mostly with brown walls in KOH. Pileocystidia absent, but scattered ± clavate to fusoid, hyaline to brownish, smooth or scatteredly diverticulate cystidioid elements of the same size as pileipellis cells present. Stipitipellis a cutis consisting of cylindrical, parallel, thick-walled (up to 1.5 μm), incrusted, up to 6.0 μm wide hyphae with brown walls in KOH. Stipe medulla hyphae similar, but ± thin-walled and hyaline to pale yellowish-brownish. Caulocystidia absent. Clamp-connections present in all tissues.– Fig. 23

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. Growing on bark of logs of Gmelina arborea.

 

Distribution. So far collected only in Zambia.

 

Revised specimens from tropical Africa.

Zambia. Kitwe, 5 Dec. 1978, G.D. Piearce FP 591 (K(M) 99655, holotype).

 

Notes. Marasmius parviconicus is characterised by having a striate pileus and a smooth and shiny stipe, and microscopically by the presence of only one type of cheilocystidia and the absence of typical pileocystidia and caulocystidia.

Similar species are especially those without developed caulocystidia. Marasmius corbariensis (Roumeg.) Singer and M. ilicis Singer differ especially by the presence of cheilocystidia in the form of broom-cells. Marasmius buxi Fr. is similar by the absence of caulocystidia and presence of lageniform to subfusoid cheilocystidia. However, it has a dark yellow-brown to red-brown pileus centre and a yellowish to almost whitish pileus margin, more distant lamellae (3–7), narrower basidiospores ((7.0–)8.5–12.5(–13) x 3.5–4.0(–4.5) μm) and well-developed pileo- and caulocystidia.

 

20. Marasmius mulanjensis Antonín

Antonín, Mycotaxon 88: 61 (2003). Type: Malawi, Mount Mulanje, 17 Nov. 1981, J. Rammeloo 7400 (BR 11970–39, holotype). 

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 61–63 (2003).

 

Pileus small, up to about 2 mm broad, convex-hemispherical to broadly conical, radially striate, (rusty) brown. Lamellae distant, L = 7–9, l = 0, narrow, ± whitish. Stipe up to 10(–15) mm long, filiform, insititious, paler at apex, black-brown towards base. (According to its photograph and exsiccatum).

 

Basidiospores 6.9–8.2 x 3.8–5.4 μm, E = 1.7–2.0, Q = 1.8, ellipsoid, subamygdaloid, thin-walled, smooth, hyaline. Basidia 19–23 x 8.5–10 μm, 4-spored, clavate. Basidioles 11–24 x 4.0–9.5 μm, clavate, subcylindrical. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, 11.5–20 x 8.5–13 μm, similar to pileipellis cells; (2) lageniform to subfusoid, 18.5–31 x 4.6–7.3 μm, rostrate, thin-walled cells, sometimes slightly thick-walled at apex. Pleurocystidia similar to cheilocystidia of type 2. Trama hyphae cylindrical, subinflated, thin-walled, mostly smooth, rarely finely incrusted, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Rotalis-type, 14–20 x 8.5–14.5 μm, (broadly) clavate, pyriform, thick-walled entirely or only in upper part, rarely entirely thin-walled; projections up to 3.0 x 1.0 μm, obtuse to acute; walls yellow-brown in KOH. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, diverticulate, up to 6.0 μm wide hyphae with dark yellow-brown to brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 24

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. Growing on dead hanging branches probably of Mumulops sp.

 

Distribution. So far collected only in Malawi.

 

Revised specimens from tropical Africa.

Malawi. Mount Mulanje, 17 Nov. 1981, J. Rammeloo 7400 (BR 11970–39, holotype).

 

Notes. Marasmius mulanjensis is characterised by having distant lamellae, rather small basidiospores, two types of cheilocystidia, well-developed pleurocystidia, a diverticulate stipitipellis and by the absence of pileo- and caulocystidia.

It differs from M. parviconicus Pegler in having a non-striate pileus, differently shaped basidiospores, two types of cheilocystidia and diverticulate stipitipellis hyphae. A similar species seems to be Marasmius corbariensis (Roumeg.) Singer from southern Europe and northern Africa. However, it has a bicoloured pileus with a darker centre and a paler margin, more numerous lamellae (L = 10–14), larger basidia (23–29 x 8.0–9.5 μm) and differently shaped basidiospores. Marasmius ilicis Singer, known from South America, has a sometimes striate pileus, only one type of cheilocystidia, and well-developed pileo- and caulocystidia (Singer 1976).

 

 21. Marasmius subalbidulus Antonín

Antonín, Mycotaxon 89(2): 428 (2004). Type: Uganda, Buganda Province, Mengo District, Mabira Forest, 9 June 1968, D.N. Pegler U 1376 (K(M) 115020, holotype). 

 

Selected descriptions and icons. Antonín, Mycotaxon 89(2): 428-430 (2004).

 

Neither macroscopic description nor slide available. Pileus 0.5–1.5 mm broad, convex to applanate, without central papilla, slightly sulcate; white, with yellowish centre. Lamellae distant, L = 7–9, l = 0–1, well-developed, shortly adnate, without collarium; white, edge entire, concolorous. Stipe up to 15 mm long, ± filiform, insititious, whitish above, brown towards base. Forming sterile stipes and rhizomorphs. (According to dry carpophores).

 

Basidiospores 8.0–9.5 x 3.5–4.5 μm, E = 1.9–2.3, Q = 2.1, pip-shaped, ellipsoid-fusoid, thin-walled, smooth, hyaline. Basidium (one found) 16 x 8.0 μm, 4-spored, broadly clavate. Basidioles 10–18 x 4.0–8.0 μm, clavate, cylindrical, often fusoid. Cheilocystidia of two types: (1) broom-cells of the Rotalis-type, 12.5–14 x 8.0–9.5 μm, clavate, pyriform, thin-walled, ± hyaline, and (2) 18–27 x 6.0–9.0 μm, lageniform to fusoid, often rostrate, obtuse to subcapitate, ± thin-walled, sometimes with scattered diverticula at basal part. Pleurocystidia in the form of cheilocystidia of type 2. Trama hyphae ± cylindrical, thin-walled, branched, smooth, hyaline, up to 8.0 μm wide. Pileipellis a hymeniderm composed of clavate, ellipsoid, pyriform to vesiculose broom-cells of the Rotalis-type, 13–21 x 9.0–17 μm, thin-walled at base, thin- to slightly thick-walled (walls up to 1.0 μm) above, warts ± narrowly conical, subacute to acute, up to 1.5 μm long; thick-walled cells subhyaline to pale yellowish-brownish in KOH. Pileocystidia 18–25 x 6.0–8.0 μm, lageniform to fusoid, often rostrate, obtuse to subcapitate, thin- to mostly slightly thick-walled, mostly with some projections in basal parts, sometimes smooth, subhyaline or with pale yellow-brown walls in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, diverticulate, up to 5.0 μm wide hyphae, with brown walls in KOH; diverticula up to 2.0 μm long, cylindrical to conical, obtuse to subacute, pale brown to dark brown in KOH. Caulocystidia scattered, 8.0–16 x 3.0–10 μm, lageniform or conical, at base with diverticula, slightly thick-walled, concolorous with stipitipellis. Clamp-connections rare, but present in all tissues. – Fig. 25

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid or amyloid.

 

Ecology. Growing on dead leaves.

 

Distribution. So far known only from Uganda.

 

Revised specimens from tropical Africa.

Uganda. Buganda Province, Mengo District, Mabira Forest, 9 June 1968, D.N. Pegler U 1376 (K(M) 115020, holotype, originally identified as M. rotalis).

 

Notes. Marasmius subalbidulus is characterised by having a white pileus, distant lamellae, well-developed ± lageniform pileo-, cheilo- and pleurocystidia, two types of cheilocystidia and small lageniform caulocystidia. It represents a very distinct species.

The only species of this section with a white pileus becoming pale yellow when dry is M. dicandidus Desjardin, Retnowati et E. Horak. It was described from Bali, Indonesia, and has larger basidiospores (8–10 x 4–5 μm) and lacks cheilo-, pleuro-, pileo- and caulocystidia (Desjardin & al. 2000). Also M. paucilamellatus Desjardin & E. Horak from Papua New Guinea is pale coloured. It especially differs by a pale yellow, then cream-tan pileus, lamellae absent when young, later distant (L = 3–5), basidiospores which are longer and narrower ((8.0–)8.5–10(–12) x 3.5–4.0 μm), cheilocystidia in the form of fusoid-mucronate cells, and absent pleuro-, pileo- and caulocystidia (Desjardin & Horak 1997).

 

Sect. Leveilleani Singer

 

Marasmius sect. Leveilleani Singer, Bull. Jard. Bot. Etat Brux. 34: 326 (1964) (as Leveilliani).

– Type species: Marasmius leveilleanus (Berk.) Pat.

 

Carpophores medium sized to rather large. Pileus usually well pigmented. Lamellae free, never collariate. Stipe central, insititious.

 

Basidiospores medium sized. Cheilocystidia present. Pleurocystidia absent. Pileipellis a hymeniderm composed of ventricose to clavate cells, smooth or often with digitate projections, transient forms present. Hyphae with clamp-connections, non-dextrinoid.

 

Notes. Singer (1964) described this section as Leveilliani derived from the original Heliomyces leveillianus. However, as the name commemorates famous botanist Joseph-Henry Léveillé, the correct spelling of the epithet is “leveilleanus”, the name of the section is Leveilleani (according to the Code, Art. 60. 11).

Pegler (1966) also placed his new species Marasmius bubalinus Pegler, described from Uganda, in this section. However, the type revision showed that it has distinctly dextrinoid hyphae and, therefore, belongs to sect. Sicci. For a detailed description and notes see there.

 

 Species description

 

Patouillard, Bull. Soc. Mycol. Fr. 33: 55 (1917). Heliomyces leveilleanus Berk., London Journ. Bot. 6: 490 (1847) (as “H. leveillianus”). Type: Sri Lanka, Hautane Range, July 1844, Gardner 72 (K(M) 99705, type).  – Marasmius umbraculum Berk. & Broome, J. Linn. Soc. Bot. 14: 36 (1873).

 

Selected descriptions and icons. Pegler, Persoonia 4(2): 118 (1966); Pegler, Kew Bull. Addit. Ser. 6: 179 (1977); Pegler, Kew Bull., Addit. Ser. 12: 154 (1986); Petch, Trans. Brit. Mycol. Soc. 31: 28–29 & Pl. IV/2 (1948); Singer, Bull. Jard. Bot. Etat Brux. 34: 326–328 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 255–256 (1965); Zoberi, Tropical Macrofungi: 74–75 (1972).

 

Pileus 7–39 mm broad, subhemispherical to convex, convex-campanulate, then applanate, umbonate, sulcate, often rugulose-sulcate, glabrous; deep reddish brown. Lamellae subclose to mostly (sub)distant, l = 0–1, subemarginate to mostly free, never collariate, thin, rather broad, not intervenose, at most slightly rugulose at base; white to yellow, with concolorous edge. Stipe 35–80 x 0.5–1.7 mm, central, (sub)cylindrical, insititious, rigid, shining, smooth, glabrous; blackish brown. Rhizomorphs present.

Context rather thick, white. (According to Singer 1964, 1965a, 1976 and Pegler 1966, 1977, 1986).

 

Basidiospores 8.5–10.5(–11.5) x 3.5–5.5 μm, E = 1.5–2.5, Q = 1.9–2.2, ellipsoid, subfusoid to subamygdaliform, thin-walled, hyaline. Basidia 25 x 7.7 μm, 4-spored, clavate. Basidioles 14–31(–34) x 3.0–8.5 μm, clavate, subcylindrical or fusoid. Cheilocystidia 14–31 x 6.9–14 μm, shaped as broom-cells of the Siccus-type, clavate to subcylindrical, slightly thick-walled in upper part, thin-walled below, projections digitate or nodulose, slightly thick-walled, up to 11 x 3.5 μm. Pleurocystidia absent. Trama hyphae cylindrical, hyaline, up to 10 μm wide; subpileipellis hyphae sometimes slightly thick-walled and brown incrusted in KOH. Pileipellis a hymeniderm composed of 15–41 x (6.0–) 8.0–15 μm, clavate to subcylindrical, sometimes irregular, slightly to distinctly thick-walled (0.5–4 μm) cells, smooth or shaped as broom-cells of the Siccus-type with transitions between them, subhyaline to yellow-brown in KOH; projections obtuse, sometimes irregular or nodulose, thick-walled, up to 7.0 x 3.1 μm. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, thick-walled (up to 2.0 μm), up to 7.5 μm wide hyphae with yellow-brown walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues.  Fig. 26

 

Chemical reactions. Neither basidiospores nor other tissues dextrinoid or amyloid.

 

Ecology. Growing on twigs, branches of bamboo and trees and pods of Leguminosae.

 

Distribution. Known from Ivory Coast, Ghana, Nigeria, Uganda and the Democratic Republic of Congo. Outside tropical Africa, it has been collected in Sri Lanka (Pegler 1986) and Mexico (Singer 1976). According to Pegler (1977) and Singer (1964), it probably represents a paleotropical species. However, Singer (1976) mentioned it from Mexico (Neotropics). Therefore, it may represent a pantropical taxon.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Yangambi, Isalowe, 26 Aug. 1938, J. Louis 11010 (BR 11480–34).

Ghana. Near Asebu, 6 June 1966, A.C. Rose B4 (K(M) 133793); Ibid., 4 June 1966, A.C. Rose A 3 (K(M) 133794).

Ivory Coast. Route de Tabou, forêt entre Troya et le fleuve Cavally (wood between Troya and the Cavally River), 27 Febr. 1975, L. Aké Assi 282 (K(M) 133795); Mount Niénokoué, 19 Jan. 1976, L. Aké Assi 445 (K(M) 133796).

Nigeria. Ibadan, Ife Biological Garden, 29 Apr. 1967, M.H. Zoberi 231 (K(M) 133797); Ibid., 28 May 1968, M.H. Zoberi 357 (K(M) 133798).

Uganda. Buganda Province, Mengo District, Mpanga Research Forest, March 1957, French 47 (K(M) 133799).

 

Revised specimens from other regions.

Sri Lanka. Hautane Range, July 1844, Gardner 72 (K(M) 99705, type; annotation added on the label: “unlocalised, ex herb. Hooker, No. 72”); Kandy District, Peradeniya, Oct. 1868, G.H.K. Thwaites 807 (K(M) 99663, syntype of M. umbraculum).

 

Notes. Marasmius leveilleanus is characterised by having a deep reddish brown pileus, a shining, smooth, glabrous and blackish brown stipe, well-developed rhizomorphs, cheilocystidia in the form of broom-cells of the Siccus-type, pileipellis cells of two transient types and non-dextrinoid hyphae.

Pegler (1977, 1986) and Singer (1964, 1965a) described smaller basidiospores (7.2–9.5 x 3.3–4.4 μm and 5.7–9.5 x 3.3–4.5 μm, respectively). Moreover, Pegler (l.c.) mentioned smaller cheilocystidia (12–15 x 4–7 μm) and smaller pileipellis cells (6–18 x 5–8 μm). In the specimens studied by me, basidiospores were constantly larger (especially broader). Zoberi (1972) described strongly intervenose lamellae, a stipe surface covered by resinaceous incrustation, smaller basidiospores (7–9 x 3–4 μm), and did not mention cheilocystidia of the Siccus-type. 

While Singer (1976, 1986) used the orthographically correct variant of the epithet “leveilleanus”, Singer (1964, 1965a), and Pegler (1966, 1977, 1986) used the spelling orthographic form “leveillianus” (see above).

 

Sect. Epiphylli Kühner

 

Marasmius sect. Epiphylli Kühner, Botaniste 25: 93 (1933) (as sect. Epiphylleae).

Type species: Marasmius epiphyllus (Pers.: Fr.) Fr.

 

Carpophores small, marasmioid. Pileus usually up to 10 mm broad, white, whitish or yellowish, membranaceous, often radially rugulose or grooved. Lamellae well-developed or reduced, sometimes absent, never collariate (false adpressed pseudocollarium sometimes present). Stipe insititious, filiform, central or eccentric, pruinose to pubescent. Rhizomorphs absent. Sterile stipes absent or present.

 

Basidiospores cylindrical, ellipsoid or lacrimoid. Basidia 2- or 4-spored. Hymenial cystidia usually present. Pileipellis a hymeniderm composed of smooth (subsect. Epiphyllini and Eufoliatini) or broom-cells (subsect. Epiphylloidei). Pileocystidia present or absent. Clamp-connections present or absent. Hyphae non-dextrinoid (subsect. Epiphyllini and Epiphylloidei) or dextrinoid (at least in stipe apex, subsect. Eufoliatini).

 

Note. Mostly temperate species, only a few species in the tropics.

Synopsis of tropical African species

Subsect. Eufoliatini Singer

 

Sect. Epiphylli Kühner, subsect. Eufoliatini Singer, Fl. Neotrop. Monogr. 17: 90 (1976).

– Type species: Marasmius eufoliatus Kühner (= M. setosus (Sowerby) Noordel.) 

Species description

23. Marasmius foliiphilus Antonín

Antonín, Mycotaxon 85: 115 (2003). Type: Cameroon, Dja Biosphere Reserve, Ekom, ca. 34 km E of Somalomo, 10 April 2001, V. Antonín Cm 01.77 (BRNM 666144, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 115–116 (2003).

 

Pileus 2–8 mm broad, low convex, with applanate, slightly obtuse to slightly depressed centre, inflexed to almost straight at the ± finely crenulate margin, glabrous, except for rugulose centre, radially striate to rugulose-striate, not hygrophanous, white. Lamellae well-developed, distant, L = 11–17, l = 0–2, slightly emarginate and broadly adnate to somewhat decurrent, not intervenose, sometimes furcate, white, with concolorous, finely pubescent edge. Stipe 5–20 x up to 0.5 mm, cylindrical, insititious, slightly broadened above, slightly broadened to subbulbose at base, entirely pubescent, hollow, entirely white when young, then white at apex and pale brown (± 6D5) towards base. Context thin, white in pileus, concolorous in stipe, without any distinct smell— Pl. 3

 

Basidiospores 6.0–8.0(–9.5) x 2.5–4.0 μm, E = 1.9–2.9, Q = 2.2–2.5, lacrimoid, ellipsoid-fusoid, thin-walled, hyaline. Basidia 18–28.5 x 6.0–8.0 μm, 4- or rarely 2-spored, clavate. Basidioles 11–24 x 3.0–7.0 μm, clavate to cylindrical. Hymenial cystidia very rare (sometimes absent?) to rather frequent, 19–33 x 6.0–11 μm, clavate, lageniform to fusoid, thin- to slightly thick-walled. Hyphae cylindrical, thin-walled, up to 8.0 μm wide. Pileipellis a hymeniderm composed of 13–42 x (3.8–)5.0–17 μm, clavate, subvesiculose, subcylindrical to subfusoid, smooth, rarely with some digitate projections or with a small rostrum, thin- to slightly thick-walled, sometimes septate cells, often in fascicles. Pileocystidia often absent or very rare, 22–50 x 5.5–9.0 μm, lageniform, subcylindrical or fusoid, thin-walled. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 5.5 μm wide hyphae with subhyaline to pale ochraceous walls in KOH; medulla hyphae up to 12 μm wide. Caulocystidia numerous, 13–85 x (4.0–)5.0–11 μm, cylindrical, clavate, lageniform, often irregular, moniliform to subcoralloid, obtuse, adpressed to erect, thin- to mostly slightly thick-walled (more distinctly at base), with subhyaline to pale ochraceous walls in KOH. Clamp-connections absent. – Fig. 27

 

Chemical reactions. Basidiospores non-dextrinoid, pileus and lamellar trama hyphae, stipe medulla and cortex hyphae weakly to distinctly dextrinoid.

 

Ecology. Single, growing in groups on dead fallen leaves and twigs, collected in forests of the Central African semideciduous type.

 

Distribution. Known from Cameroon, the Democratic Republic of Congo, Nigeria and Uganda.

 

Revised specimens from tropical Africa.

Cameroon. Dja Biosphere Reserve, Ekom, ca. 34 km E of Somalomo, 10 April 2001, V. Antonín Cm 01.77 (BRNM 666144, holotype); South West Province, Korup National Park, trail to Mana Bridge, 12 April 1997, P.J. Roberts K 1134 (K(M) 91513, BRNM 691106).

Democratic Republic of Congo. Tshopo Province, Ngene–Ngene, 18 April 1984, B. Buyck 1473 (BR 11750–13).

Nigeria. ? Cross–River State, Calabar-Cameroon road, 23 June 1990, R.A. Nicholson (K(M) 18620, as M. lolema).

Uganda. Masaka District, Bukoto County, Kasonko forest near Kiwala, 31 May 1971, K. Arnstein Lye M 90 (K(M) 111895, as M. lolema).

 

Notes. Marasmius foliiphilus is characterised by having entirely white carpophores (except for the basal part of the stipe when mature), well-developed lamellae, small basidiospores, often absent (sometimes rare) pileocystidia, rare to frequent hymenial cystidia, clampless and dextrinoid hyphae. Having smooth pileipellis cells and dextrinoid hyphae, it belongs to the subsect. Eufoliatini Singer.

Species belonging to this subsection are very rare in the tropics. Singer (1965b, 1976) described two South-American species without clamps, M. sanctixaverii Singer and M. caliensis Singer. Both of them differ especially in having larger basidiospores (8.2–9.7 x 3.8–4.5 μm and 6.0–10.0 x 2.5–3.5 μm, respectively)

 

Sect. Fusicystides Singer

 

Marasmius sect. Fusicystides Singer, in Singer & Digilio, Lilloa 25: 287 (1952).

Type species: Marasmius fusicystis Singer (= M. isabellinus Pat.)

 

Carpophores marasmioid. Pileus pigmented. Lamellae not collariate. Stipe non-insititious, oblique, eccentric to lateral, sometimes rudimentary. Rhizomorphs absent.

 

Basidiospores large. Cystidia conspicuous, sometimes incrusted. Pileipellis never hymeniform, but a cutis consisting of the Ramealis-structure or with irregularly arranged broom-cells (sometimes similar to the genus Marasmiellus). Hyphae, at least in some tissues, dextrinoid. Clamp-connections present.

 

Notes. This section is macroscopically and microscopically very similar to the genus Marasmiellus. The most important differential features are the non-insititious stipe and particularly, the dextrinoid hyphae.

Species of this section have not been recorded in tropical Africa so far.

 

Species description

 

24. Marasmius longicystidiatus Antonín

Antonín, Mycotaxon 85: 119 (2003). Type: Malawi, Nyika National Park, Gîte Chelinda, 6 Dec. 1981, J. Rammeloo 7703 (BR 11995–64, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 119–121 (2003).

 

Pileus up to ± 10 mm broad, convex to plano-convex, finely tomentose, radially rugulose to sulcate, pinkish ochraceous to ochraceous. Lamellae distant, L = 6–13, l = 0–1, ± free to adnate, ± pliciform when young, then well-developed, not branched, not intervenose, pale orange-ochraceous. Stipe well-developed, eccentric to sublateral, up to 3 x 1 mm, ± cylindrical to laterally compressed, curved, non-insititious, tomentose to hairy, whitish to pale ochraceous, with small whitish basal disc. [Notes according to a photograph.] — Pl. 3

 

Basidiospores (14–)14.5–16.0(–17.5) x 5.4–6.6(–7.0) μm, E = 2.2–2.9, Q = 2.5, clavate or lacrimoid, thin-walled, hyaline. Basidia 37–50 x 8.1–11.5 μm, 4-spored, clavate. Basidioles 15–44 x 3.5–10 μm, clavate, subcylindrical, subfusoid. Hymenial cystidia 45–115 x 7.0–14 μm, fusoid, subacute, thin-walled. Trama hyphae ± cylindrical, ± thin-walled, up to 10 μm wide. Pileipellis a cutis, in some part transient to a (sub)hymeniderm, composed of three types of structures: (1) radially arranged, interwoven, ± cylindrical to subinflated, branched, smooth or incrusted, up to 8.0 μm wide hyphae with pale orange-ochraceous walls in KOH; hyphae sometimes (scatteredly) diverticulate or with lateral projections (slightly developed Ramealis-structure); (2) clavate, subcylindrical to subfusoid, regular to irregular, lobate, ± thin-walled, smooth, ± hyaline cells; (3) broom-cells of the Siccus-type, 16–25 x 6.0–13 μm, clavate or subcylindrical, regular or irregular, entirely or at least in upper part slightly thick-walled, with pale orange-ochraceous walls in KOH, with irregular, nodulose to subcoralloid, obtuse, slightly thick-walled projections. Pileocystidia absent. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae. Caulocystidia 23–39 x (3.8–)6.2–9.2 μm, ± cylindrical, conical, subfusoid, sublageniform, irregular, often branched to subcoralloid, slightly thick-walled. Clamp-connections present. – Fig. 28

 

Chemical reactions. Basidiospores non-dextrinoid; hyphae dextrinoid.

 

Ecology. Growing on dead twigs in a montane forest (2300 m alt.).

 

Distribution. Known only from the type locality in Malawi.

 

Revised specimens from tropical Africa.

Malawi. Nyika National Park, Gîte Chelinda, 6 Dec. 1981, J. Rammeloo 7703 (BR 11995–64, holotype).

 

Notes. Marasmius longicystidiatus is characterised by having a sulcate, pinkish ochraceous to ochraceous pileus, distant, pale orange-ochraceous lamellae, large basidiospores, very long basidia and hymenial cystidia, and a pileipellis composed of three types of elements without pileocystidia. Only two other species from this section are known so far (Singer 1986). The neotropical species M. isabellinus Pat. (= M. fusicystis Singer), known from Argentina, Bolivia, Brazil and Ecuador, has narrower basidiospores (4.0–5.8 μm), smaller basidia (28–36 x 7.5–9.0 μm), smaller cystidia (20–80 x 4.0–10.0 μm), and well-developed pileocystidia (Singer 1976); the paleotropical M. campanella Holterm. (=? M. rufescens Berk. & Broome) has, according to Singer (1976), narrower basidiospores (x 4.3–4.5 μm).

 

Sect. Chordales Fr.

 

Marasmius sect. Chordales Fr., Epicrisis: 381 (1838).

Marasmius sect. Alliacei Kühner, Botaniste 25: 87 (1933) (as Alliateae).

Marasmius, I Longipedes Morgan, J. Mycol. 11: 237 (1905).

Type species: Marasmius chordalis Fr.

 

Carpophores moderately large, marasmioid or collybioid. Pileus up to 50 mm broad, often hygrophanous, often slightly rugulose. Lamellae well-developed, thin, distant to moderately close, free or adnate, sometimes attached to a pseudocollarium. Stipe central, non-insititious, usually with a distinct basal mycelium. Rhizomorphs absent.

 

Basidiospores ellipsoid, amygdaliform, limoniform or clavate. Basidia 2- or 4-spored. Cheilocystidia usually present, sometimes absent. Pleurocystidia present or absent. Pileipellis hymeniform and composed of smooth cells, rarely cells with some digitate projections. Pileocystidia absent or less frequently present. Hyphae never dextrinoid. Caulocystidia present or absent.

 

Notes. Singer (1964, 1965a) also included Marasmius ferrugineoluteus Beeli (Bull. Soc. Roy. Bot. Belg. 60: 155. 1928) in this section. I revised the type specimen (Democratic Republic of Congo, Eala, Ipeko, May 1923, M. Goossens–Fontana 142, BR 11449–03, holotype) with the following results: Basidiospores 6.9–9.2 x 3.3–4.6 μm, ellipsoid, thin-walled. Basidia 21.5–25.5 x 5.4–6.9 μm, 4-spored, clavate. Basidioles 14–25.5 x 4.0–7.7 μm, clavate, subcylindrical, subfusoid. Cheilocystidia 21.5–35 x 3.8–6.9 μm, subcylindrical to clavate, with irregular to coralloid projections on top, thin-walled. Pleurocystidia 23–29 x 5.4–8.0 μm, fusoid, subutriform, sometimes subcapitate. Hyphae cylindrical, hyaline or slightly brownish, thin-walled, seemingly slightly gelatinised, up to 8.0 μm wide. Pileipellis a hymeniderm composed of 21–31 x 6.9–14 μm, clavate, broadly clavate, pyriform, thin- to slightly thick-walled cells, with (sub)hyaline to dark brown walls (especially at base) in KOH. Subpileipellis composed of cylindrical, thick-walled, irregular, branched, strongly gelatinised, smooth or incrusted, up to 9.5 μm wide hyphae with subhyaline to brown walls in KOH. Pileocystidia 23–101 x 5.0–6.2 μm, cylindrical, sublageniform or subulate, obtuse, often irregular at base, thick-walled, brown in KOH (more intensively towards base), sometimes incrusted at base. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 4.5 μm wide hyphae with yellow-brown to brown walls in KOH. Stipe surface covered by long cylindrical, thick-walled (up to 2.0 μm), 2.5–6.2 μm wide, obtuse to subacute, sometimes irregular (especially at base), sometimes branched, yellow-brown to brown hairs. Clamp-connections present in all tissues. Neither basidiospores nor other structures dextrinoid.

Singer (1964) already mentioned that this species may belong to a different genus (Flammulina) especially based on the colour of carpophores, the ± velutinose stipe surface, and gelatinised hyphae. I fully agree with Singer´s opinion, and have excluded it from the genus Marasmius.

 

Key to tropical African species

 

1. Basidiospores small, up to 6.5 μm long and 3.5 μm broad; pileus white, yellowish, purple or lilac grey to cinereous ................. 2

1*. Basidiospores larger, (6.0–)8.0–11.2 x 3.1–5.2 μm; pileus buff, pale grey-brown to beige coloured ............... 3

 

2. Pileus and stipe dark purple; cheilo- and caulocystidia present; pileipellis cells and stipitipellis hyphae with brownish-purplish content ....................... 26. M. pegleri

2*. Pileus white-yellow, never dark purple; cheilocystidia and caulocystidia absent; pileipellis cells and stipitipellis hyphae hyaline ............ 25. M. lolema

 

3. Basidiospores (8.5–)9.0–11.2 x 3.1–4.0(–4.4) μm; pileus pale grey-brown to beige coloured; stipe eccentric, 18–22 mm long; pleurocystidia present; caulocystidia 19–80 x 3.8–14 μm ................ 27. M. schreursii

3*. Basidiospores 6.0–10.0 x 4.0–5.2 μm; buff cream coloured; stipe central, about 35 x 3.5 mm; pleurocystidia absent; caulocystidia 13–35 x 7.0–15 μm ................................................  28. M. mvumae

 

Species descriptions

 

25. Marasmius lolema Beeli

Beeli, Bull. Soc. Roy. Bot. Belg. 60: 154 (1928). Type: Democratic Republic of Congo, Equateur Province, Eala, Sept. 1923, M. Goossens–Fontana 298 (BR 11486–40, holotype).

 

Selected descriptions and icons. Beeli, Bull. Soc. Roy. Bot. Belg. 60: 154 (1928); Singer, Bull. Jard. Bot. Etat Brux. 34: 341–342 (1964); Singer, Flore Icon. Champ. Congo, fasc. 14: 261 & Pl. 45, fig. 4 (1965).

 

Pileus 25–35 mm broad, convex or plano-convex, often inflexed, thin and rugulose-crenulate at margin, concentrically sulcate, glabrous, white-yellow. Lamellae shortly adnate to almost free, extremely crowded and narrow, white. Stipe 70–80 x 2.5–3 mm, cylindrical or subcylindrical, hollow, glabrous, smooth, white, with some yellowish zones or stains (according to Beeli: white-yellow, except for white base); basal mycelium abundant, tomentose, white. Context not changing colour, white, fibrose, thin at margin, subcartilaginous in stipe. (According to Beeli 1928a and Singer 1964). — Pl. 3

 

Basidiospores 3.1–4.6 x 2.5–3.1 μm, E = 1.2–1.6, Q = 1.4, broadly ellipsoid, ± thin-walled, hyaline. Basidia 17–21 x 4.2–5.8 μm, 4-spored, clavate. Basidioles 15–23.5 x 3.5–8.5 μm, clavate or subcylindrical. Cheilocystidia not found; some irregular, submonilifirm basidioles found on lamellar edge. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, up to 17 μm wide. Pileipellis a hymeniderm or subhymeniderm composed of 24–61 x 8.5–20 μm, clavate, subfusoid, lageniform, sometimes irregular or moniliform, ± thin-walled cells. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 7 μm wide hyphae. Caulocystidia absent; some scattered clavate to cylindrical, adpressed to suberect, obtuse cells present. Clamp-connections present in all tissues. – Fig. 29

 

Chemical reactions. Neither spores nor other structures dextrinoid.

 

Ecology. On dead leaves in an inundated forest.

 

Distribution. Known from Uganda, Tanzania and the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Equateur Province, Eala, Sept. 1923, Goossens–Fontana 298 (BR 11486–40, holotype).

Tanzania. Eastern Province, Kilosa District, Ilonga, Matarawe River, 25 May 1968, D.N. Pegler T 1070 (K(M) 134246).

 

Notes. Marasmius lolema is characterised by having a very rich basal tomentum, very small spores, an often subhymeniform (weakly hymeniform) pileipellis and by the absence of any cystidia. 

Only one so far known species, M. pegleri Courtec. (= M. purpureus Pegler) described from Tanzania, has similar small spores (3.0–4.5 x 2.7–3.5 μm). However, it differs especially by its dark purple pileus and stipe, broad, brown lamellae, and by the presence of cheilo- and caulocystidia.

However, having a subhymeniform pileipellis, Marasmius lolema may represent a transition to some other genus (Hydropus?).

 

26. Marasmius pegleri Courtec.

Courtecuisse, Doc. Mycol. 14 (54–55): 89 (1984). Type: Tanzania, Southern Highlands Province, Iringa, Kiban, Mufindi, Imgoda Tea Estate, 11 May 1968, D.N. Pegler T 905 (K(M) 99657, holotype). – Marasmius purpureus Pegler, Kew Bull. Addit. Ser. 6: 167 (1977); non M. purpureus Berk. & M.A.Curtis, J. Linn. Soc. Bot. 10: 299 (1868).

 

Selected descriptions and icons. Courtecuisse, Doc. Mycol. 14(54–55): 87–89 (1984); Pegler, Kew Bull. Addit. Ser. 6: 167 (1977).

 

Pileus 4–6 mm broad, conico-campanulate, umbonate, surface uniformly dark purple, glabrous, smooth, striate. Lamellae sinuate-adnexed, very pale brown, fairly broad, distant, with occasional lamellulae. Stipe 35–40 x 0.5–1 mm, cylindrical, hollow, surface concolorous with the pileus, finely pruinose towards the apex, arising from a white basal mycelium. Context thin, pale brown. (According to Pegler 1977).

 

Basidiospores 3.0–4.5 x 2.7–3.5 μm, E = 1.1–1.5, Q = 1.3, broadly ellipsoid to subglobose, smooth, thin- to very slightly thick-walled, smooth, hyaline, with a refractive vacuole, often in tetrads in preparations. Basidia 18–20 x 6.0–7.5 μm, 4-spored, clavate. Basidioles 12–20 x 3.0–7.0 μm, cylindrical, clavate. Cheilocystidia 15–38 x 4.5–12 μm, clavate, subcylindrical, (sub)utriform, thin-walled, hyaline. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, up to 15 μm wide, with hyaline walls. Pileipellis a hymeniderm composed of 17–36 x 8.0–16 μm, clavate, broadly clavate, smooth, sometimes subcapitate cells, with subhyaline to pale greyish-brownish-purplish contents in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, ± slightly thick-walled, up to 5.0 μm wide hyphae, with pale greyish-purplish walls in KOH. Caulocystidia 23–49 x 6.0–10 μm, single or in groups, clavate, cylindrical, subfusoid, sometimes capitate, thin- to slightly thick-walled, obtuse. Clamp-connections present in all tissues.  Fig. 30

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. Growing among dead leaves on a forest floor.

 

Distribution. It has been collected only in the type-locality in Tanzania.

 

Revised specimens from tropical Africa.

Tanzania. Southern Highlands Province, Iringa, Kiban, Mufindi, Imgoda Tea Estate, 11 May 1968, D.N. Pegler T 905 (K(M) 99657, holotype).

 

Notes. Marasmius pegleri is characterised by having a uniformly dark purple coloured pileus and stipe, very small basidiospores and brownish-purplish coloured medulla hyphae.

For differences with M. lolema also having very small spores see above. Marasmius ionides Pat. found in Guadeloupe also has a violaceous pileus. However, it has a “rufescent” or whitish to pale brown stipe, larger basidiospores (5.5–7.2 x 3.2–3.8 μm) and hair-like subulate caulocystidia (600–700 x 10–19 μm) which are thick-walled (Pegler 1983; Singer 1976).

 

27. Marasmius schreursii Antonín

Antonín, Mycotaxon 88: 64 (2003). Type: Democratic Republic of Congo, Katanga Province, Luiswishi, 6 Febr. 1986, J. Schreurs 1038 (BR 7881–24, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 64–66 (2003).

 

Pileus 22 mm broad, convex to plano-convex, obtuse or with a low broad papilla at centre, slightly striate at margin, smooth, hygrophanous, pale grey-brown to beige coloured, more distinctly grey when wet. Lamellae distant, L = ± 15, l = 2–3, broadly adnate to emarginate with a small tooth, rather broad, ventricose, whitish or greyish, edge concolorous or yellowish (when becoming dry ?). Stipe slightly to distinctly eccentric, 18–22 x 1.5–2.5 mm, cylindrical, broadened above, tomentose, whitish, pale greyish towards base. (Notes according to a photograph and an exsiccatum). — Pl. 3

 

Basidiospores (8.5–)9.0–11.2 x 3.1–4.0(–4.4) μm, E = 2.1–3.2, Q = 2.7, narrowly ellipsoid, cylindrical-ellipsoid to sublacrimoid, thin-walled, hyaline. Basidia 27–34.5 x 5.8–8.0 μm, 4-spored, clavate. Basidioles 15–35 x 3.3–7.0 μm, clavate, cylindrical or subfusoid. Cheilocystidia 21.5–34.5 x 10–19 μm, broadly clavate, pyriform, sometimes capitate, thin- or slightly thick-walled. Pleurocystidia 20–40 x 4.0–6.2 μm, narrowly clavate, narrowly fusoid, sublageniform, often (sub)rostrate, thin-walled. Trama hyphae cylindrical to subinflated, thin- to slightly thick-walled, up to 16 μm wide, mixed with narrow thick-walled ones. Pileipellis a hymeniderm composed of 10–27 x 6.2–19 μm, clavate, pyriform to vesiculose, slightly to distinctly thick-walled, often irregular, pale greyish-brownish in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 7.0 μm wide hyphae. Caulocystidia 19–80 x 3.8–14 μm, cylindrical, narrowly clavate, subfusoid, sublageniform, sometimes irregular or furcate, slightly thick-walled. Clamp-connections present in all tissues. – Fig. 31

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. On dead wood in a “forêt dense”.

 

Distribution. Known only from the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Katanga Province, Luiswishi, 6 Febr. 1986, J. Schreurs 1038 (BR 7881–24, holotype).

 

Notes. Marasmius schreursii is characterised by having narrowly ellipsoid, cylindrical-ellipsoid basidiospores and by the presence of cheilo-, pleuro- and caulocystidia.

A close species seems to be M. notandus Corner described from Borneo (Corner 1996). In comparison with Corner´s description, it differs only in having an umber brown pileus, robust (up to 40 x 8 mm) and at base bulbose stipe, not more than a finely white pruinose stipe surface, longer pleurocystidia (50–70 x 7–10 μm) and clampless hyphae. Marasmius batistae Singer, known from Brazil (Pegler 1997; Singer 1976), with a similar pileus colour, has smaller basidiospores (5.5–8.8 x 2.5–3.8 μm), smaller basidia (18–25 x 4.3–5.0 μm), hyaline pileipellis cells and lacks cystidia; all species with cystidia described by Singer (1976) have smaller or broader basidiospores.

 

28. Marasmius mvumae Antonín & C. Sharp

Antonín & C. Sharp in Antonín, Mycotaxon 88: 63 (2003). Type: Zimbabwe, Midland Province, Mvuma, Beacon Hill Plots, 1930 A4, 9 Jan. 1997, C. Sharp 1213/99 (BR 152505–21, holotype). 

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 63–64 (2003).

 

Pileus about 20 mm broad, plano-convex, with regular or irregular, sometimes crenulate margin, matt, buff cream coloured. Lamellae shortly adnate, distant, firm, white. Stipe about 35 x 3.5 mm, cylindrical, central, firm, matt, buff coloured at apex, fulvous and then sienna towards base. Context strongly acrid.

 

Basidiospores 6.0–10.0 x 4.0–5.2 μm, E = 1.5–2.1, Q = 1.7–1.9, ellipsoid, ± thin-walled, smooth, hyaline; thick-walled spores (crassospores) also present. Basidia 24–31 x 8.0–9.0 μm, 4-spored, clavate. Basidioles 15–33 x 3.5–9.0 μm, clavate to cylindrical. Cheilocystidia 13–26 x 9.0–15 μm, clavate, pyriform or subfusoid, thin-walled, hyaline. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, smooth, up to 12 μm wide. Pileipellis a hymeniderm composed of 15–27 x 11–17 μm, clavate, pyriform, lageniform, thin-walled cells. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 8.0 μm wide hyphae. Caulocystidia 13–35 x 7.0–15 μm, adpressed to erect, clavate to cylindrical, obtuse, thin- to slightly thick-walled. Clamp-connections present in all tissues. – Fig. 32

 

Chemical reactions. Neither basidiospores nor other structures dextrinoid.

 

Ecology. Caespitose, on thick leaf litter in a miombo woodland, under Brachystegia glaucescens.

 

Distribution. Known only from the type locality in Zimbabwe.

 

Revised specimens from tropical Africa.

Zimbabwe. Midland Province, Mvuma, Beacon Hill Plots, 1930 A4, 29 Dec. 1996, C. Sharp 483/96 (BR 152504–20); Ibid., 9 Jan. 1997, C. Sharp 1213/99 (BR 152505–21, holotype).

 

Notes. Marasmius mvumae is characterised by having ± buff coloured caespitose carpophores, rather small basidiospores, well-developed cheilo- and caulocystidia and by the absence of pileo- and pleurocystidia.

Among similar species, M. amabilis Hariot & Pat., with about the same size and colour of carpophores, has distinctly larger basidiospores (e.g. 26.5 x 6.5 μm) (Desjardin & Horak 1997), M. oligocystis Singer from Brazil is smaller (pileus 6 mm broad), with small basidiospores (5.3–6 x 2.3–3 μm), M. aimara Singer from Bolivia has a pale golden cinnamon coloured pileus and stipe and smaller basidiospores (5.4–5.6 x 4.3 μm) (Singer 1976).

 

Sect. Neosessiles Singer

 

Marasmius sect. Neosessiles Singer, Mycologia 50: 104 (1958).

– Type species: Marasmius neosessilis Singer

 

Carpophores pleurotoid. Pileus small, well-pigmented or not. Lamellae not or indistinctly collariate. Stipe absent or rudimentary, and then often eccentric or lateral, insititious, subinsititious or with basal mycelium.

 

Basidiospores moderately large to large. Pileipellis a hymeniderm or loose hymeniderm composed of broom-cells of the Siccus-type. Hyphae dextrinoid (at least in stipe) or non-dextrinoid, mostly with clamp-connections or rarely without them.

 

Note. Species of this section are very close to sect. Sicci, and may even belong there. They differ only in having pleurotoid carpophores and a rudimentary or absent stipe.

Key to tropical African species

1. Basidiospores 15–19 x 4.5–6.0 μm, fusoid, clavate; pleurocystidia present, numerous; pileus orange brown ................................ 32. M. aff. cecropiae

1*. Basidiospores smaller, ellipsoid or cylindrical-ellipsoid; pleurocystidia present (but sometimes indistinct) or absent; pileus never orange brown ................................... 2

 

2. Lamellae absent or only very narrow and vein-like (mostly two); basidiospores 8.2–11 x (4.0–)4.5–5.5 μm, ellipsoid to cylindrical-ellipsoid; pleurocystidia absent ............................................. 33. M. cyphella

2*. Lamellae well-developed; basidiospores smaller or larger; pleurocystidia present (but sometimes indistinct) or absent ...... 3

 

3. Basidiospores small, 6.9–8.5 x 4.6–5.4 μm, ellipsoid; basidia 23–27 x 7.7–8.1 μm; cheilocystidia 11.5–16.5 x 5.4–7.7 μm; stipitipellis hyphae diverticulate or with broom-cells of the Siccus-type on the stipe surface; pleurocystidia present but indistinct.............................................................. 31. M. cf. sejunctus

3*. Basidiospores larger, 8.0–12.5 x 4.0–7.0(–8.0) μm, ellipsoid, broadly ellipsoid, subamygdaliform or ellipsoid-fusoid; basidia broader, up to 11(–13) μm broad; cheilocystidia larger, 12–21 x 9.0–13 μm or 15.5–27.5 x 7.7–10 μm, respectively; pleurocystidia present or absent ................................................... 4

 

4. Pileus off-white to light beige, beige brown-grey to greyish-yellow with a tinge to yellowish white, covered by dark brown to almost black, somewhat spiny granules; lamellae not intervenose to slightly intervenose, sometimes branched (old carpophores); basidiospores 9.0–12.5(–13.1) x 5.0–7.0(–8.0) μm, ellipsoid to broadly ellipsoid; basidia 24–34.5 x 9.0–13.0 μm; cheilocystidia 15.5–27.5 x 7.7–10 μm; pleurocystidia indistinct to distinct, 30–48 x (6.0–)7.5–13 μm ................................ 30. M. bururiensis

4*. Pileus almost white at first, then salmon flesh coloured, pinkish cinnamon or sordid salmon, without granular covering; lamellae intervenose when old; basidiospores 8.0–11(–12) x 4.0–5.5(–6.0) μm, ellipsoid, subamygdaliform, ellipsoid-fusoid; basidia up to 27 x 11 μm; cheilocystidia 12–21 x 9.0–13 μm; pleurocystidia absent ......... 29. M. neosessilis

 

Species descriptions

 

29. Marasmius neosessilis Singer

Singer, Mycologia 50: 103 (1958). Type: Argentina, Iguazú, Cataratas, Singer & Digilio 57 (LIL, holotype, not studied).

 

Selected descriptions and icons. Morris, Kirkia 13(2): 341 (1990); Nicholson, Nigerian Field 54: 25 (1989); Pegler, Kew Bull. Addit. Ser. 6: 202–204 (1977); Singer, Fl. Neotrop. Monogr. 17: 261–263 (1976).

 

Pileus 1–19 mm broad, reniform in outline, glabrous when young, then cross-veined and radially deeply sulcate-grooved when mature, at first almost white but soon “Apricot” to salmon flesh (9F7), pinkish cinnamon (11B8) or sordid salmon (12B10). Lamellae distant, L = 3–4 (in larger caps ± 6), l = 1–5, adnate or in smaller caps decurrent, not intervenose and not intermixed in small carpophores, then with numerous veins, narrow, white. Stipe absent in small carpopohores even at maturity, or present, 0.5–2 x 0.1–1 mm, rudimentary and adpressed to both substratum and carpophore, slightly subtomentose, insititious, apex whitish, below pallid to pale fuscous, later fuligineous. Context thin, white, inodorous. (According to Singer 1976).

 

Basidiospores 8.0–11(–12) x 4.0–5.5(–6.0) μm, E = 1.7–2.5, Q = 1.9–2.1, ellipsoid, subamygdaliform, ellipsoid-fusoid, thin-walled, smooth. Basidia e.g. 24 x 11 μm, 4-spored, clavate. Basidioles 10–27 x 5.0–11 μm, clavate, cylindrical or subfusoid. Cheilocystidia in the form of broom-cells, 12–21 x 9.0–13 μm, clavate, subcylindrical, thin-walled, hyaline; projections ± thin-walled. Pleurocystidia absent. Trama hyphae cylindrical or subinflated, ± thin-walled, hyaline, smooth to minutely incrusted, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–25(–37) x 7.0–14(–20) μm, (broadly) clavate, subcylindrical, sometimes branched, thin-walled with slightly thick-walled apex, with 8–20(–25) short, up to 5.0 x 1.5 μm, obtuse, digitate to conical, slightly thick-walled projections; all thick-walled parts with reddish yellow walls in KOH. Clamp-connections present in all tissues.  Fig. 33

 

Chemical reactions. Basidiospores non-dextrinoid, hyphae dextrinoid (at least some of them and weakly).

 

Ecology. Single or in groups, growing on dead twigs.

 

Distribution. It was described from Argentina, and is known also from Ecuador and Argentina in South America. In tropical Africa, it has been found in Ghana, Ivory Coast, Kenya, Nigeria, and Uganda.

 

Revised specimens from tropical Africa.

Ghana. Cape Coast, near Kakomdo, 25 March 1967, A.C. Rose CC 6712 (K(M) 115017).

Ivory Coast. Forêt de la Besso, 31 March 1975, L. Aké Assi 329 (K(M) 115016).

Kenya. Western Province, Kakamega District, Kakamega Forest, near Forest Station, 20 Jan. 1970, L. Ryvarden (K(M) 5438); ? Nyanza Province, Kericho District, Kericho, Kigumu River, 25 March 1968, D.N. Pegler K 238 (K(M) 11523).

Nigeria. Cross River State, Calabar-Ikar Road, 17 July 1990, R.A. Nicholson 640 (K(M) 18826).

Uganda. Buganda Province, Mengo District, Mabira Forest, 9 June 1968, D.N. Pegler U 1380 (K(M) 115024).

 

Notes. Marasmius neosessilis is characterised by having ± pleurotoid carpophores with absent or rudimentary stipe, interveined lamellae when old, moderately large basidiospores, dextrinoid hyphae and lacking pleurocystidia.

In comparison with the microscopic description by Singer (1976), all collections mentioned here have broader basidiospores; they are only 3.3–4.0 μm broad according to Singer. In my opinion, the slightly broader (and also slightly longer) basidiospores measured in this material fall within the variability of this species. Pegler (1977) mentioned slightly broader basidiospores than Singer (7–10 x 3.5–4.5 μm), and his description also differs by narrower cheilocystidia (12.5–17 x 5–9 μm) and smaller pileipellis broom-cells (7–12 x 4.5–11 μm).

Collection Pegler K 238 (Kenya, K(M) 115023) differs from other African specimens in having distinctly larger basidiospores (10–13(–15) x 5.5–8.0 μm). Other features were identical. Therefore, it is included with a question mark here.

 

30. Marasmius bururiensis Antonín

Antonín, Mycotaxon 85: 111 (2003). Type: Burundi, Bururi Province, Siguyaye Valley, W from the road Mutambara–Bururi, 3 Febr. 1979, J. Rammeloo 6509 (BR 11926–92, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 111–113 (2003).

 

Pileus 5–10 mm broad, shell-shaped, strongly convex, involute, thin, distinctly sulcate-striate, very pale off-white to light beige, beige brown-grey to greyish-yellow (± 4B3) with a tinge of yellowish white (4A2), striae slightly paler; covered by dark brown to almost black, somewhat spiny granules, largest and closest to each other at point of attachment. Lamellae distant, L = 6–10, l = 0(–1), adnate, rather thick, 1–1.5 mm broad, not intervenose to slightly intervenose, sometimes branched (old carpophores), slightly convex, off-white, edge whitish, granulose. Stipe very short to absent, eccentric to lateral, ca. 3 x 1 mm, ± cylindrical, strongly curved, very shortly adpressed fibrillose, yellowish white above, brownish-greyish to brown at base. Context without a special smell— Pl. 3

 

Basidiospores 9.0–12.5(–13.1) x 5.0–7.0(–8.0) μm, E = 1.4–2.1, Q = 1.7–1.8, ellipsoid to broadly ellipsoid, hyaline, thin-walled. Basidia 24–34.5 x 9.0–13 μm, 4-spored, clavate. Basidioles 15.5–31.5 x 2.5–12.5 μm, clavate, cylindrical, subfusoid. Cheilocystidia in the form of broom-cells, 15.5–27.5 x 7.7–10 μm, clavate to broadly clavate, entirely thin-walled, rarely slightly thick-walled in upper part, (sub)hyaline; projections irregular, ± cylindrical, nodulose or coralloid, slightly thick-walled, up to 11.5 x 3.8 μm; similar but smooth cells rarely present among broom-cell cheilocystidia. Pleurocystidia distinct or indistinct, 30–48 x (6.0–)7.5–13 μm, fusoid, often rostrate, obtuse, rarely subacute, thin-walled, not incrusted, sometimes seemingly slightly mucronate at apex. Trama hyphae cylindrical, smooth, ± hyaline, up to 10 μm wide, slightly brown pigmented or incrusted in subpileipellis; often slightly thick-walled, often inflated near septum, up to 10(–16) μm wide in stipe medulla. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 9.2–23 x (5.4–)7.7–15.5 μm, clavate to vesiculose, less frequently subcylindrical, slightly to distinctly thick-walled (especially in upper part), basal part sometimes thin-walled, hyaline to brown in KOH; projections digitate, obtuse to subacute, often irregular or nodulose, up to 7.5 x 2.5 μm. Stipitipellis a cutis consisting of parallel, slightly thick-walled, smooth or incrusted, up to 7.0 μm wide hyphae. Caulocystidia (terminal cells) adpressed to often (sub)erect, often forming groups of clavate, cylindrical or subfusoid, thin- to slightly thick-walled, obtuse, up to 12 μm wide cells. Clamp-connections present in all tissues. – Fig. 34

 

Chemical reactions. Stipitipellis and stipe medulla hyphae weakly dextrinoid, other structures non-dextrinoid.

 

Ecology. Gregarious, growing on dead twigs in a montane rainforest and a gallery forest.

 

Distribution. Known from Burundi, Cameroon and Tanzania.

 

Revised specimens from tropical Africa.

Burundi. Muramvya Province, Teza, 22 Dec. 1978, J. Rammeloo 6241 (BR 11916–82); Bururi Province, Siguyaye Valley, W from the road Mutambara-Bururi, 3 Feb. 1979, J. Rammeloo 6509 (BR 11926–92, holotype).

Cameroon. South–West Province, Korup Forest Reserve, Mundemba, 27 Jan. 1989, R. Watling s.n. (E).

Tanzania. West Usambara Mts., Shume-Magamba Forest Reserve, E slope of Mabweni, 14 Febr. 1985, I. Krisai 3749 (WU).

 

Notes. Marasmius bururiensis is characterised by having a rather beige brown-grey to greyish-yellow, sulcate and spiny-granulose pileus, distant lamellae, a very short to absent stipe, rather broad basidiospores, and by the presence of cheilo- and pleurocystidia. 

As for similar species, M. griseoroseus (Mont.) Dennis, from South America (Bolivia, Colombia, Panama), has a different pileus colour and larger and narrower basidiospores ((9.5–)11.5–14.5 x 2.2–4.0 μm according to Singer 1976; 13–15.5 x 3.5–4.0 μm according to Dennis 1970); M. ustilago Singer, from Bolivia, has a deeply fuscous pileus, a distinct stipe, narrower basidiospores (9.5–11 x 5.3–5.5 μm), and two types of context hyphae (Singer 1976); M. cecropiae Dennis, from South America (Bolivia, Venezuela), has a pale ochraceous, on drying fulvous orange to orange-buff coloured pileus, longer and narrower basidiospores (11–14 x 4.0–4.8 μm) and fusoid cheilocystidia (Dennis 1961; Singer 1976); M. paulensis Singer, from Brazil, has an orange to pale orange pileus with concolorous lamellae, larger fusoid basidiospores (12.5–15 x 4.5–6.2 μm), filamentous-cylindrical, rarely clavate pleurocystidia, rarely with some terminal projections, and a subhymeniform pileipellis (Singer 1976). Of species with indistinct or absent pleurocystidia, M. spaniophyllus Berk. var. spaniophyllus, collected in Brazil, has a brown pileus, non-intervenose lamellae and no pleurocystidia; M. spaniophyllus var. iguazuensis (Singer) Singer, described from Argentina and known also from Venezuela, has a white, then pale to greyish fuscous pileus and no pleurocystidia. Marasmius neosessilis Singer has a different pileus colour, smaller basidiospores (8.0–11(–12) x 4.0–5.5(–6.0) μm), smaller basidia and cheilocystidia, and lacks pleurocystidia, and Marasmius sessilis (Pat.) Sacc. & P. Syd., known from tropical Asia, differs especially by the absence of cystidia and smaller basidiospores (6–8 x 4–5 μm) (Patouillard 1896).           

 

31. Marasmius cf. sejunctus Singer

Marasmius sejunctus Singer, Fl. Neotrop. Monogr. 17: 260 (1976).

 

Macroscopical description not available, material sparse. According to the exsiccatum, it seems that pileus was up to 2 mm broad, not white and the lamellae were not intervenose or ramified.

 

Basidiospores 6.9–8.5 x 4.6–5.4 μm, E = 1.4–1.7, Q = 1.6, ellipsoid, thin-walled, hyaline (only a few basidiospores found). Basidia 23–27 x 7.7–8.1 μm, 4-spored, clavate. Basidioles 11.5–24.5 x 3.5–9.2 μm, clavate, subcylindrical or subfusoid. Cheilocystidia in the form of broom-cells of the Siccus-type, 11.5–16.5 x 5.4–7.7 μm, clavate to subcylindrical, thin- or slightly thick-walled above, hyaline. Pleurocystidia scattered, slightly projecting beyond hymenium, 28–30 x 7.0–7.5 μm, fusoid, often subrostrate, obtuse. Trama hyphae cylindrical, thin-walled, smooth, up to 7.5 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, clavate, rarely subcylindrical, thick-walled entirely or only at apex, subhyaline to brown in KOH; projections digitate, often irregular or (sub)coralloid, sometimes branched, obtuse, thick-walled, up to 8.0 x 2.5 μm. Subpileipellis of branched, cylindrical or subinflated, slightly thick-walled, hyaline to brownish, up to 8.0 μm wide hyphae. Stipitipellis a cutis consisting of parallel, cylindrical, thick-walled, smooth or incrusted, often brown pigmented, 3.5–7.5 μm wide hyphae. Caulocystidia in the form of wart-like projections (diverticulate surface hyphae) or typical broom-cells of the Siccus-type, up to 20 x 9.2 μm, clavate, subcylindrical or subfusoid, thick-walled, brown in KOH; projections irregular to nodulose, thick-walled, obtuse. Clamp-connections present. – Fig. 35

 

Chemical reactions. Stipe medulla hyphae weakly dextrinoid, other stuctures non-dextrinoid.

 

Ecology. Growing on twigs, in a tropical virgin forest with Macrolobium dewevrei.

 

Distribution. Known only from the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Yambao, ca. 20 km north–east of the village, 22 June 1939, J. Louis 15288 (BR 11498–52).

 

Notes. Collection Louis 15288 is identified as M. neosessilis on the box label in the herbarium BR. However, it differs from M. neosessilis especially in having non-dextrinoid hyphae in the pileus context. The exact identification of this collection is not possible because of the absence of a macrodescription. Microscopically the closest species seems to be M. sejunctus Singer, found in Venezuela and Bolivia. Its microscopic description (Singer 1976) agrees rather well with our fungus. Other similar species are M. tenuissimus (Jungh.) Singer, which differs by all hyphae non-dextrinoid and larger basidiospores (8.0–10.5 x 3.5–4.5 μm) (Pegler 1997; Singer 1976), and M. sessiliaffinis Singer [= M. spaniophyllus Berk. var. sessiliaffinis (Singer) Singer] with absent pleurocystidioid elements and larger pileipellis cells with a different type of projections.

 

32. Marasmius aff. cecropiae Dennis

Marasmius cecropiae Dennis, Kew Bull. 15: 92 (1961).

 

Macrodescription not available. Small orange brown sessile carpophores with up to ca. 3 mm broad pileus and sparse lamellae (dry specimens).

 

Basidiospores 15–19 x 4.5–6.0 μm, E = 3.1–4.0, Q = 3.5, fusoid, clavate, thin-walled, smooth, non-dextrinoid. Basidioles 10–26 x 4.0–8.0 μm, clavate, cylindrical, fusoid. Cheilocystidia in the form of broom-cells, 10–22 x 5.0–11 μm, clavate, subcylindrical, thin-walled, hyaline, some cells almost coralloid and branched. Pleurocystidia numerous, 25–35 x 8.0–10 μm, clavate, fusoid, thin-walled, hyaline. Trama hyphae ± cylindrical, thin-walled, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–17 x 8.0–12 μm, clavate, subcylindrical, thin-walled, with up to 5.0 x 1.0 μm large, obtuse to subacute, nodulose, slightly thick-walled projections; thick-walled parts reddish yellowish in KOH. Clamp-connections present in all tissues. – Fig. 36

 

Chemical reactions. Neither basidiospores nor hyphae dextrinoid.

 

Ecology. Gregarious, growing on suspended twigs in troops.

 

Distribution. Collected only in Ghana.

 

Revised specimens from tropical Africa.

Ghana. Cape Coast, near Kekomdo, on suspended twigs in troops, 23 May 1970, A.C. Rose CC 7004A (K(M) 108852, as M. griseoroseus); Ibid., near Jukwa, 2 June 1970, A.C. Rose CC 7004B (K(M) 111246, as M. griseoroseus).

 

Notes. This fungus is characterised by having large basidiospores, well-developed pleurocystidia and non-dextrinoid hyphae.

These specimens were originally identified as M. griseoroseus. I revised a duplicate (?) of the holotype specimen of M. griseoroseus (Mont.) Dennis (French Guyana, on rotten bamboo, Leprieur 691, K(M) 108877), however, the material is very poor, and I did not find any basidiospores. According to the dry specimens, our fungus is macroscopically really similar to M. griseoroseus. However, both Singer (1976) and Pegler (1983) described smaller basidiospores: Singer (9.5–)11.5–14.5 x 2.2–4 μm and Pegler 11–16 x 3.5–4.5 μm. Moreover, pleurocystidia are not mentioned in descriptions.

According to the descriptions (Dennis 1961; Singer 1976), M. cecropiae seems to be similar to our fungus in macroscopic and some microscopic features. However, its basidiospores are smaller (11–14 x 4–4.8 μm), some hyphae are weakly dextrinoid and it grows on fallen dicotyledonous leaves (always of Cecropia?). Because of the absence of a macroscopic description, I have decided to publish this species as M. aff. cecropiae.

 

33. Marasmius cyphella Dennis & D.A. Reid

Dennis & D.A. Reid, Kew Bull. 1957/2: 288 (1957). Type: Malaysia, on leaves and twigs of Hevea, R.N. Hilton s.n. (K(M)108854, holotype).

 

Selected descriptions and icons. Corner, Nova Hedw. Beih. 111: 45 (1996); Dennis & Reid, Kew Bull. 1957/2: 288 (1957).

 

Pileus 1–5 mm broad, thin-membranaceous, reniform, non-lamellate or with low vein-like lamellae (mostly 2) and attached to substrate by rudimentary lateral stipe. In dried specimens the hymenial surface is cream coloured and the upper surface olivaceous brown. [According to Dennis & Reid 1957 and collector´s notes]

 

Basidiospores 8.2–11 x (4.0–)4.5–5.5 μm, E = 1.6–2.3, Q = 1.9–2.1, ellipsoid, cylindrical-ellipsoid, ellipsoid-fusoid, thin-walled, smooth. Basidia 23–38 x 7.5–9.0 μm, 4-spored, clavate. Basidioles 15–45 x 3.0–10 μm, narrowly clavate or cylindrical. Cheilocystidia in the form of broom-cells, 13–22 x 4.0–9.0 μm, (narrowly) clavate, cylindrical, thin- to slightly thick-walled, hyaline. Pleurocystidia absent. Trama hyphae ± cylindrical, thin-walled, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 10–18 x 7.0–12 μm, (broadly) clavate, vesiculose, often branched, thin- to slightly thick-walled, with short and wide, up to 3.0(–4.0) x 1.5(–2.0) μm, obtuse, not nodulose, thin- to slightly thick-walled projections; scattered smooth cells present. Clamp-connections present in all tissues. – Fig. 37

 

Chemical reactions. Basidiospores non-dextrinoid, hyphae non-dextrinoid or dextrinoid.

 

Ecology. Gregarious, growing on fallen dead leaves and on old living twigs and leaves.

 

Distribution. Described from Malaya. In tropical Africa, it has been found in Ghana.

 

Revised specimens from tropical Africa.

Ghana. Ashanti, on leaves, June 1923, H.K. Hewison (K(M)108879).

 

Revised specimens from other regions.

Malaysia. On leaves and twigs of Hevea, R.N. Hilton (K(M)108854, holotype).

 

Notes. Marasmius cyphella is characterised by having small carpophores without lamellae or with only vein-like lamellae, moderately large, ellipsoid, cylindrical-ellipsoid, ellipsoid-fusoid basidiospores, cheilocystidia in the form of broom-cells and by the absence of pleurocystidia. According to Dennis & Reid (1957) it may cause thread-blight of Hevea in Malaysia.

Dennis & Reid (1957) mentioned a collection from Ghana (originally identified as Marasmius scandens and later changed to M. cyphella), which should differ from M. cyphella by having two low vein-like lamellae, slightly larger basidiospores and more numerous pileipellis broom-cells. However, they proposed to include it in the variability of. M. cyphella and not to describe it as a separate species, with which I agree.

Marasmius ustilago Singer, described from Bolivia, has a ± similar size of basidiospores, however, it has a deep fuscous pileus and stipe; M. sessiliaffinis Singer, from Panama and Bolivia, has slightly smaller basidiospores (7.8–9.7 x 4.7–5.5 μm) and a white to fuscous or grey pileus; M. tenuissimus (Jungh.) Singer, known from Brazil, Bolivia and Indonesia, has a larger rusty brown to greyish fuscous pileus (7–14 mm broad) and narrower basidiospores (8–10.5 x 3.8–4.7 μm); M. spaniophyllus Berk., from South America, has larger basidiospores (9–12.3 x 4.3–6.5 μm) and a white to pale argillaceous, pale fuscous or dirty brown pileus, and M. sejunctus Singer, from Bolivia and Venezuela, has a salmon pinkish to pinkish ochraceous pileus (Singer 1976).

 

Sect. Globulares Kühner

 

Marasmius sect. Globulares Kühner, Botaniste 25: 100 (1933) (ut Globularinae). 

Type species: Marasmius globularis Fr. (= M. wynnei Berk. & Broome)

 

Carpophores marasmioid or collybioid. Pileus smooth or sulcate, white or mostly pigmented. Lamellae well-developed, adnate to free. Stipe central, non-insititious, with well-developed basal mycelium.

 

Basidiospores small to very large, smooth, non-dextrinoid, inamyloid; thick-walled, dextrinoid spores present in some species. Pileipellis a hymeniderm composed of smooth cells. Cheilocystidia present. Pleurocystidia present or absent, if present then often with refractive contents. Caulocystidia present or absent. Hyphae dextrinoid. Clamp-connections present.

 

Notes. Species with setose cystidia sometimes included in this section really belong to sect. Sicci, series Spinulosi (Clémençon) Desjardin. Generally, characters of species of sect. Globulares are almost identical with those of species of sect. Sicci. They differ only in having smooth pileipellis cells (those of Sicci possessing broom cells). All other features support the opinion that these sections (Globulares and Sicci) are identical. I have separated both sections based on the tradition of recent monographs.

Marasmius jodocodos Henn. was included in this section by Pegler (1977) based on his collections from Uganda. However, I revised 7 specimens identified as M. jodocodos preserved in the Kew herbarium and found all of them having a non-hymeniform pileipellis and distinctly amyloid basidiospores. Therefore, Marasmius jodocodos s. Pegler belongs to the genus Mycena. I consider the name M. jodocodos s. orig. a nomen dubium (see chapter “Exluded and dubious names”). Also M. brunneodiscus Pegler (Kew Bull. 21: 525 (1968); K(M) 92582!, holotype) has a pileipellis composed of chains of (sub)globose to ellipsoid cells and amyloid basidiospores. Therefore, also this species belongs to the genus Mycena.

 

 Key to tropical African species

 

1. Pleurocystidia absent .......................................................................................... 2

1*. Pleurocystidia present  ................................................................................ 14

 

2. Basidiospores shorter than 11.5 μm ................................................................ 3

2*. Basidiospores longer than 11.5 μm ........................................................... 7

 

3. Basidiospores shorter than 7.5 μm ....................................................... 4

3*. Basidiospores longer, 6.2–11.5 μm ............................................................ 5

 

4.  Pileus pale cream; lamellae not intervenose; basidiospores 5.8–6.6 x 3.1–3.7 μm; cheilocystidia 15.5–23 x 3.5–5.4 μm, subcylindrical, lageniform; caulocystidia 11.5–23 x 6.2–7.9 μm ........ 37. M. kigwenensis

4*. Pileus lilac grey to cinereous; lamellae strongly intervenose; basidiospores (5.2–)6.0–7.0(–7.5) x 3.5–4.5 μm; cheilocystidia 10–30 x 7.5–15 μm, (broadly) clavate, subfusoid, pyriform; caulocystidia 20–48 x 10–22 μm .......  38. M. favoloides

 

5. Pileus not sulcate, white, milky white or cream, sometimes yellow-orange coloured at centre; basidiospores up to 4.5 μm broad; caulocystidia always numerous; growing on decaying wood and twigs .................................................... 6

5*. Pileus slightly sulcate, pale ochraceous with ochraceous-brown margin or white with yellowish brown centre; basidiospores 7.0–11.5 x 3.8–5.4(–6.6) μm; caulocystidia absent or present; growing on dead leaves .........................  36. M. albertianus

 

6. Stipe white or cream at apex, red-brown towards base, densely fasciculate, especially in lower part, 50–170 x 1–2 mm; pileus white, whitish or cream coloured, with yellow-orange or brownish tinge at centre; basidiospores (6.9–)8.1–11.5 x 3.0–4.0(–4.2) μm, Q = 2.4–2.8; cheilocystidia 16.5–31 x 6.6–12(–15.5) μm; pileipellis cells 19–35(–46) x (11–)13–19(–23) μm; caulocystidia (19–)24–54(–77) x 7.7–12.5 μm .........................................  34. M. arborescens

6*. Stipe entirely white to cream when young, later brown towards base, single or in small groups; pileus uniformly yellowish white; basidiospores 6.2–8.5(–10.0) x 3.5–4.5 μm, Q = 1.8–2.2, cheilocystidia (11.5–)15–25 x 3.5–6.9 μm; pileipellis cells 17.5–27 x (7.0–)11.5–15(–17) μm; caulocystidia 19–35(–44) x (3.8–)5.4–7.7 μm ............................. 35. M. lacteoides

 

7. Pileus violaceous or lilac coloured or at least with such tinge ........................................... 8

7*. Pileus without violaceous or lilac tinge ........................................................................ 12

 

8.  Pileus rather small, 21–40 mm broad, violaceous, violaceous brown, brown with violaceous tinge or pale vinaceous with ochraceous centre .............................................................................. 9

8*. Pileus larger, 30–70(–100) mm broad, whitish to ochraceous, with brown-violaceous centre and then non-striped or pale and dark violaceous or violaceous and then distinctly radially striped ................... 10

 

9. Pileus violaceous, violaceous brown or brown with violaceous tinge, striped; stipe violaceous above, brown towards base; basidiospores 15.5–22.5 x 3.3–5.0 μm ..................................... 39. M. violaceoides

9*. Pileus pale violaceous with ochraceous centre, not striped; stipe 140–150 x 1.5–4 mm, yellowish brown towards apex, darker brown at base; basidiospores 10.5–16.5(–15.5) x 4.5–5.3(–6.7) μm .............................. 40. M. mesosporus

 

10. Pileus whitish to ochraceous, with brown-violaceous centre, non-striped; basidiospores large, (21–)23–28(–30) x 5.5–6.5(–7.0) μm ..................................................................................... 55. M. camerunensis

10*. Pileus pale and dark violaceous or violaceous and then yellow striped, spores smaller and/or narrower .................. 11

 

11. Pileus large, 30–100 mm broad, rugulose at centre, lilac pinkish to dark lilac-violaceous, pale or pale violaceous on sulci, without yellow tinge; lamellae pale lilac; stipe (40–)120–170 x 2–12 mm; context mild with slightly acrid after-taste ................... 41. M. zenkeri

11*. Pileus smaller, 30–67(–100) mm broad, violaceous and lemon yellow to yellow striped; lamellae yellowish white or pale lemon yellow, without violaceous tinge; stipe 80–150 x 2.5–6(–10) mm; context with bitter taste ....... 42. M. bekolacongoli

 

12. Basidiospores 15.5–25.5 x 3.8–5.4 μm; pileus fuligineous brown or ochraceous, often with whitish striae (striped); lamellulae present; stipe 70–180 x 3–6 mm; context with acrid taste .............................. 43. M. brunneolus

12*. Basidiospores longer and broader, 20–28 x 5.0–7.0 μm; pileus differently coloured – ochraceous to yellow-ochraceous or rusty; lamellulae absent; stipe smaller, up to 120 mm long; context with mild taste (or taste unknown) ................... 13

 

13. Pileus ochraceous at centre, yellow ochraceous on sulci, pallescent up to cream towards margin in striae, striation less distinct; stipe 85 x 2.5 mm; cheilocystidia (12.5–)15.5–28.8 x 6.2–11.5(–21) μm, variable in shape, clavate, subvesiculose, cylindrical, sometimes irregular, lobed or subrostrate, rarely subcoralloid .....................................  44. M. tshopoensis

13*. Pileus rusty coloured; stipe 120 x 5–6 mm; cheilocystidia 15–18 x 9.0–12 μm, clavate .................  45. M. missangoënsis

 

14. Basidiospores shorter than 10 μm ............................................................... 15

14*. Basidiospores longer than 10 μm ................................................. 17

 

15. Pileus small (8–18 mm broad), umbonate; stipe densely fasciculate; caulocystidia absent ......................  46. M. witteanus

15*. Pileus larger (15–70 mm broad), never distinctly umbonate; stipe never densely fasciculate ................ 16

 

16. Pileus distinctly striate at margin, cream, with fuligineous brown to ochraceous centre and often differently coloured striae; stipe subglabrous; basidiospores 6.2–7.9(–9.2) x 3.5–4.2; cheilocystidia 24–31(–38.5) x 10–14 μm, clavate, ± regular; pleurocystidia 48–100 x 12.5–20(–23) μm, clavate to fusoid; caulocystidia absent ..................... 47. M. goossensiae

16*. Pileus not striate, brown beige at centre, paler, greyish orange, towards margin; stipe entirely mealy to finely pubescent; basidiospores 5.0–6.2 x 2.9–3.7 μm; cheilocystidia 10.0–23(–35) x 4.0–7.0 μm, clavate, subcylindrical, subfusoid, often irregular; pleurocystidia 19–55 x 8.5–17 μm, variable, cylindrical, clavate, fusoid; caulocystidia present ............................................  48. M. muramwyanensis

 

17. Carpophores robust, collybioid, similar to M. oreades; pileus 6–80 mm, never sulcate, never radially striped, reddish brown at centre, yellow-orange towards margin; stipe 27–60 x 4–12 mm, finely tomentose, white to whitish above, orange-cream to brownish orange towards base; basidiospores 10.8–14.5 x (4.2–)4.6–6.0(–6.6) μm, ellipsoid to subfusoid; cheilocystidia large, (16–)22–61.5 x 6.2–12.5 μm, variable; pleurocystidia 27–46.5 x 5.5–11.5 μm, clavate, subfusoid; caulocystidia present, 15.5–42 x (4.6–)5.1–11.0 μm .......................................................  50. M. heinemannianus

17*. Carpophores less robust (but they can be very large), marasmioid; pileus sulcate, often radially striped; stipe less robust (up to 4 mm wide); spores clavate to subfusoid; caulocystidia absent or present ........ 18

 

18. Pileus without violaceous tinge, not radially striped ..................................... 19

18*. Pileus violaceous coloured, radially striped ......................................................... 22

 

19. Basidiospores longer than 23 μm; cheilocystidia 12–23 μm long ............................................ 20

19*. Basidiospores 13.5–15.5 x 3.1–3.5 μm; cheilocystidia longer than 26 μm ............................................. 21

 

20. Pileus 20–30 mm broad, yellowish; stipe 1–1.5 mm wide; caulocystidia present ........... 51. M. flavidulus

20*. Pileus up to 100 mm broad, beige cream coloured with darker centre; stipe up to 8 mm wide; caulocystidia absent ................................ 52. M. latepileatus

21. Pileus yellow to dirty yellow; stipe 80 x 2–2.5 mm, brown-red; basidiospores 13.5–15 x 3.1–3.5 μm; cheilocystidia 28.5–46 x 13–18.5 μm; pleurocystidia 40–77 x 11.5–20(–23) μm ................. 49. M. flavus

21*. Pileus whitish to pale cream at centre, almost white when dried out, yellowish grey towards margin; stipe 50–90 x 0.5–1.2 mm, whitish or cream at apex, orange brown to (orange) golden at base; basidiospores 16.5–23 × 3.5–5.0 μm; cheilocystidia 10–31 × 6.0–11 μm; pleurocystidia (21–)27–48 × (4.5–)6.5–12 μm ............  53. M. albidocremeus

 

22.  Basidiospores 16.0–28.5 x (4.5–)5.5–6.0 μm, Q = 3.7–4.7; cheilocystidia 14–26 x 7.5–11 μm, clavate, subfusoid, vesiculose ...................  54.1. M. staudtii var. staudtii

22*. Basidiospores 32–44 x 5.5–7.5 μm, E = 4.8–6.8, Q = 5.9; cheilocystidia 24–35(–42.5) x (8.5–)10–13.5(–17) μm ...............................  54.2. M. staudtii var. magnisporus

Species descriptions

34. Marasmius arborescens (Henn.) Beeli

Beeli, Bull. Soc. Roy. Bot. Belg. 60: 156 (1928). Collybia arborescens Henn., Bot. Jahrb. Syst. 22: 106 (1895). Type: Cameroon, Bipinde, 1898, in: Zenker, Flora von Kamerun, No. 1362 (BR 11380–31, as Collybia arborescens, neotype). – Mycena fasciculata Beeli, Bull. Soc. Roy. Bot. Belg. 59: 82 (1927).

 

Selected descriptions and icons. Heim, Ann. Sci. Nat., sér. Bot., 9: 1–8 (1948); Pegler, Kew Bull. Addit. Ser. 6: 173–175 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 350–351 (1964); Singer, Flore Icon. Champ. Congo 14: 265–266 (1965); Zoberi, Tropical macrofungi: 76–79 (1972).

 

Pileus 10–20(–28) mm broad, (sub)campanulate or conical-subglobose with involute margin, then convex to applanate with slightly depressed centre and slightly inflexed margin, dry, glabrous, slightly hygrophanous, slightly translucently striate towards margin when young, almost up to ½ of diam. when old, smooth or slightly rugulose, white or whitish or with cream (3A2), yellow-orange or brownish tinge at centre. Lamellae close, L = 28–32, l = 2–4, adnexed to adnexed-subdecurrent, narrow (up to 1 mm), pale yellowish (4A2) with pale cream orange reflex or whitish, with concolorous, finely pubescent, entire to slightly uneven edge. Stipe 45–170 x 0.7–2 mm, (sub)cylindrical, often compressed, mealy, pubescent to tomentose, hollow, white or concolorous with pileus at apex, dirty reddish brown, beige-brown, ochraceous-brown to dark red-brown towards base (± 6D5–6C5, 7D–E7); stipes densely fasciculate, especially in lower part, non-insititious; with cream, tomentose basal mycelium. Context thin (up to ± 0.5 mm), white or whitish, rather fragile when fresh and dry, concolorous with surface in stipe, with a smell of bitter almonds (like Marasmius oreades, Heim 1948) or fungoid (Rammeloo) and mild, pleasant, fungoid taste— Pl. 4

 

Basidiospores (6.9–)8.0–11.5 x 3.0–4.0(–4.2) μm, E = (2.0–)2.2–3.0(–3.8), Q = 2.4–2.8, narrowly ellipsoid to sublacrimoid, hyaline; often forming thick-walled, smooth, dextrinoid chlamydospores inside. Basidia 17–24 x 6.2–8.5 μm, 4-spored, clavate. Basidioles 11–27.5 x 3.8–9.0 μm, clavate, subfusoid, subcylindrical. Cheilocystidia 16.5–40 x 6.6–15 μm, clavate, subfusoid, subvesiculose, sometimes slightly irregular, sometimes subrostrate, rarely with one projection, thin-walled. Pleurocystidia absent. Trama hyphae ± cylindrical, thin-walled, branched, up to 15 μm wide. Pileipellis a hymeniderm composed of 19–35(–46) x (11–)13–19(–23) μm, clavate to (sub)vesiculose, thin-, rarely slightly thick-walled cells, hyaline in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, thin- to slightly thick-walled, up to 7 μm wide hyphae. Caulocystidia numerous, (19–)24–54(–77) x 7.7–15 μm, cylindrical, clavate, subfusoid, sometimes irregular, rarely subcapitate, thin-walled. Clamp-connections present in all tissues. – Fig. 38

 

Chemical reactions. Basidiospores (except for chlamydospores), hymenium and pileipellis cells non-dextrinoid; hyphae and caulocystidia dextrinoid.

 

Ecology. On stumps and decaying wood, e.g. of Ceiba pentandra; noted from Brachystegia woodland and riparian evergreen forest (Morris 1990), a primary forest of the Central African semideciduous type and Cocoa-tree plantations.

 

Distribution. Widely distributed throughout tropical Africa. It is known from Angola, Burundi, Cameroon, the Democratic Republic of Congo, Ghana, Kenya, Malawi (Morris 1990), Nigeria, Tanzania and Uganda.

 

Revised specimens from tropical Africa.

Angola. Golungo Alto near Banga, April 1856, Welwitsch 295 (K(M) 134278).

Burundi. Bururi Province, Kigwena, Kigwena Forest, 20 Feb. 1979, J. Rammeloo 6668 (BR 13427–41); Ibid., 22 Feb. 1979, J. Rammeloo 6710 (BR 11946–15).

Cameroon. Bipinde, 1898 & 1899, in: Zenker, Flora von Kamerun, No. 1362 (BR 11380–31, PC (s.n.), K(M) 92583, as Collybia arborescens, duplicates of neotype); Ebolowa, 16 Aug. 1946, R. Heim s.n. (PC); N´Kongsamba, plaine de Lelem, 22 July 1946, R. Heim s.n. (PC); Dja Biosphere Reserve, Somalomo, ca. 14 km ESE of the village, 9 April 2001, V. Antonín Cm 01.58 (BRNM 666131); Mbalmayo Forest Reserve, Oyack II village, 20 Sept. 2002, leg. C. Douanla–Meli (HUYI).

Democratic Republic of Congo. Nlemfu, 6 May 1907, Van Tilborg s.n. (BR 11385–36, as Collybia arborescens); Urselia, 2 April 1923, Ghesquière 45 (BR 11382–33); Ipamu, Oct. 1921, Vanderyst 11079 (BR 11384–35); Kalo, 1928, M. Goossens–Fontana 409 (BR 11381–32); Equateur Province, Eala, Sept. 1923, M. Goossens–Fontana 229 (BR 11378–29, holotype of Mycena fasciculata).

Ghana. Tafo, 29 April 1957, M. Holden GC 157 (K(M) 134276).

Kenya. Western Province, Kakamega Forest, ca. 13 km ESE of Kakamega, 25–27 Febr. 1973, L. Ryvarden 9550 (K(M) 134272).

Nigeria. Ibadan, Ife Biological Garden, 1967, M.H. Zoberi 114 (K(M) 134268); Ibadan, University of Ife, Botanic Garden, May 1963, S.O. Alasoadura (K(M) 134269); Ife, Oct. 1967, C.T. Ingold (K(M) 134271); Ibadan, Nigerian College of AST, 7 July 1957, D.J. Hankler 19 (K(M) 134270); Cross River State, Calabar-Cameroon Road, Oban Forest, 16 Aug. 1990, R.A. Nicholson 720 (K(M) 16551).

Tanzania. Tanga Province, Muheza District, Amani, 9 Apr. 1971, H. Faulkner 4539 (K(M) 134279).

Uganda. Mpanga Baseline, Makerere College, 16 April 1964, E.A. Calder 48 (K(M) 134274); Ibid., 20 April 1964, E.A. Calder 64 (K(M) 134275); Buganda Province, Mengo District, Zika Forest, N of Entebbe, 12 June 1968, D.N. Pegler U 1436 (K(M) 134277); Masaka District, Bukoto County, SW end of Kako Forest, 5 May 1972, K. Arnstein Lye M 139 (K(M) 134273).

 

Notes. Having small densely fasciculate carpophores with a white to whitish pileus, Marasmius arborescens belongs to the most distinctive species of this section. Moreover, it has rather small and narrow basidiospores, clavate, subfusoid or subvesiculose cheilocystidia and numerous, (19–)24–54(–77) x 7.7–12.5 μm large, cylindrical, clavate or subfusoid caulocystidia.

In comparison with the literature, Singer (1964, 1965a) mentioned narrower basidiospores (x 3.0–3.5 μm), smaller basidia (13–14 x 5.7 μm), and smaller caulocystidia (25–40 x 7.0–7.5 μm), Pegler (1977) also mentioned narrower basidiospores (x 3.0–3.5 μm), smaller basidia (12.5–18 x 4.0–5.0 μm), smaller cheilocystidia (13–22 x 9.0–11 μm), and smaller caulocystidia (15–40 x 4.0–12 μm). The description published here agrees (except for the narrower basidiospores, x 3.0–3.5 μm) with a very detailed description by Heim (1948) based on findings from Cameroon. He also tested positively the presence of hydrocyanic acid using the Guignard´s method. Heim (1948) described var. alba (invalidly, without a Latin diagnosis) which differs by a slightly larger and white, only at centre ochraceous pileus, and white-cream coloured lamellae. He also proposed the generic name Sympodia for it (Heim 1948: 8).

No similar species with so densely fasciculate growth has been described so far. A densely cespitose stipe (but no so distinct) is also present in M. witteanus Singer, which differs by a ± ochraceous brown pileus, smaller basidiospores [(4.6–)5.0–6.2(–6.6) x (2.5–)3.1–3.8(–4.2) μm], larger refractive cheilocystidia and well-developed pleurocystidia.

Mycena fasciculata Beeli is mentioned as a synonym of this species by Pegler (1977). However, two specimens are marked as “type” in the BR herbarium. The first specimen (“holotype”, Goossens–Fontana 229, BR 11378–29 !) really represents Marasmius arborescens, whereas the second specimen marked as “co-type” (Goossens-Fontana 265, BR 11379–30 !) collected in the same locality represents a fungus belonging to sect. Sicci. Therefore, Mycena fasciculata really represents a synonym of M. arborescens.

According to Walleyn & Rammeloo (1994), M. arborescens belongs to fungi used as food in the Democratic Republic of Congo. It is known under the local name “mobwati” (Hendrickx 1948). 

 

35. Marasmius lacteoides Antonín

Antonín, Mycotaxon 85: 118 (2003). Type: Zambia, Chowo Forest, 12 Dec. 1981, J. Rammeloo 7856 (BR 12032–04, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 118–119 (2003).

 

Pileus 8–30 mm broad, broadly conical-hemispherical when young, then convex, obtuse or with small broad papilla at centre, with involute, then straight and undulate, only slightly translucently striate margin, hygrophanous, smooth, glabrous, later rugulose, whitish yellowish, whitish when dried out. Lamellae close, L = ca. 60, with irregularly arranged lamellulae, emarginate with tooth, narrow, white to cream. Stipe 35–60 x 2–3 mm, cylindrical, slightly broadened above, slightly clavate at base, finely pubescent, entirely milky white when young, then brownish towards base, whitish at apex and through pale brown to chestnut brown towards base when old. Context white, concolorous with surface under pellis, hollow in stipe. — Pl. 4

 

Basidiospores 6.2–8.5(–10.0) x 3.5–4.5 μm, E = 1.5–2.7, Q = 1.8–2.2, ellipsoid to sublacrimoid, hyaline. Basidia 18.5–26 x 6.5–8.5 μm, 4-spored, clavate. Basidioles 10.0–24 x 3.3–8.0 μm, clavate, fusoid, cylindrical. Cheilocystidia (11.5–)15–25 x 3.5–6.9 μm, clavate, subfusoid, subcylindrical, often irregular, sometimes lobed or branched, thin-walled. Pleurocystidia absent. Hyphae of cylindrical, subfusoid or subinflated, ± thin-walled, hyaline, up to 15 μm wide cells, mixed (in pileus) with thick-walled, 1.5–19 μm wide hyphae. Pileipellis a hymeniderm composed of 17.5–27 x (7.0–)11.5–15(–17) μm, clavate to pyriform, thin- to slightly thick-walled cells, smooth, hyaline in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, thin- to slightly thick-walled, up to 5.0 μm wide hyphae, with ± hyaline walls in KOH. Caulocystidia numerous, 19–35(–44) x (3.8–)5.4–7.7 μm, adpressed to erect, cylindrical, narrowly clavate, lageniform, sometimes irregular or branched, thin- to slightly thick-walled. Clamp-connections present in all tissues.  Fig. 39

 

Chemical reactions. Basidiospores, hymenium and pileipellis cells non-dextrinoid; hyphae and caulocystidia dextrinoid.

 

Ecology. On rotting wood and dead twigs in litter, e.g. of Brachystegia.

 

Distribution. So far known from Zambia, and probably also from Kenya and Malawi.

 

Revised specimens from tropical Africa.

Kenya. Western Province, Kakamega Forest, ca. 13 km ESE of Kakamega, 25–27 Jan. 1973, L. Ryvarden 9449 (K(M) 8420, as M. arborescens).

Malawi. ? Makwawa, Zomba, 6 March 1980, B. Morris 154 (K(M) 8421, as M. arborescens).

Zambia. Chowo Forest, 12 Dec. 1981, J. Rammeloo 7856 (BR 12032–04, holotype); Ibid., 10 Dec. 1981, J. Rammeloo 7821 (BR 12023–92).

 

Notes. Marasmius lacteoides is characterised by having milky white carpophores, very close lamellae, small basidiospores, no pleurocystidia and well-developed caulocystidia. The collection J. Rammeloo 7821 slightly differs from the type collection in having smaller and slightly differently shaped cheilocystidia (more distinctly clavate, (10.0–)11.5–19 x 4.6–6.9 μm), and on average slightly smaller pileipellis cells (13–19 x 7.0–13 μm).

Macroscopically, it is rather similar to M. arborescens (Henn.) Beeli which differs by a densely fasciculate stipe, coloured red-brown towards base, slightly longer and narrower basidiospores with different Q-ratio [(6.9–)8.1–11.5 x 3.0–4.0(–4.2) μm, Q = 2.4–2.8], larger cheilocystidia (16.5–31 x 6.6–12(–15.5) μm), larger pileipellis cells [19–35(–46) x (11–)13–19(–23) μm], and larger caulocystidia [(19–)24–54(–77) x 7.7–12.5 μm].

Marasmius niveus Mont., from South America, differs by a larger (30–50 mm), strongly sulcate pileus, subclose to distant lamellae, a glabrous stipe with a fulvous to chestnut base, narrower basidiospores (5.3–9.0 x 2.7–3.5 μm, but 8.0–11 x 2.5–3.0 μm according to Dennis 1970) and by the absence of caulocystidia (Singer 1976); M. strictipes (Peck) Singer, from Americas, has a larger (26–60 mm) pileus with a reticulose-rugose centre, a large stipe (47–81 x 1.5–5.5(–8) mm) with an orange-cinnamomeous base, and narrow (26–28 x 2.0–5.0 μm), filamentous, cylindrical-flexuose to subclavate cheilocystidia; M. pellucidus Berk. & Broome, from Sri Lanka and New Caledonia, has a reddish brown stipe towards base, distant lamellae (L = 12–20), smaller basidiospores (6.0–7.5 x 2.5–3.5 μm), larger obtusely fusoid cheilocystidia (15–40 x 5–7 μm) and only thin-walled caulocystidia (Desjardin & Horak 1997), however, according to Pegler (1986) the basidiospores are 8–10.5 x 3.0–4.0 μm.

  

36. Marasmius albertianus Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 351 (1964). Type: Democratic Republic of Congo, Lacs Édouard et Kivu District, Albert National Park, Kalonge s. Butahu, 8 May 1953, de Witte 8969 (BR 11375–26, holotype).

 

Selected descriptions and icons. Pegler, Kew Bull. Addit. Ser. 6: 174 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 351–352 (1964); Singer, Flore Icon. Champ. Congo 14: 266 (1965).

 

Pileus about 20 mm broad, convex, umbilicate, slightly sulcate, glabrous, pale ochraceous with ochraceous-brown zone towards margin, darker at centre. Lamellae close, L = ca. 35–40, adnexed to emarginate-adnexed, rather narrow, brownish when dry. Stipe 70–80 x 1.5–3 mm, subcylindrical, smooth, (sub)glabrous, chestnut subfuligineous; with scattered pale basal mycelium. Context thin. (According to Singer 1964, 1965a).

 

Basidiospores 7.0–11.5 x 3.8–5.4(–6.6) μm, E = 1.4–2.2, Q = 1.7–1.9, (broadly) ellipsoid to subamygdaliform, thin-walled. Basidia 22.5–23 x 6.9–7.7 μm, 4-spored, clavate. Basidioles 11.5–25 x 4.0–7.0 μm, cylindrical, clavate, subfusoid. Cheilocystidia 21–36 x 5.0–11 μm, clavate, subfusoid, subcylindrical. Pleurocystidia absent. Trama hyphae ± cylindrical, ± thin-walled, hyaline, up to 15.5 μm wide; mixed with ± cylindrical, sometimes irregular, thick-walled, hyaline, 2.5–23 μm wide hyphae. Pileipellis a hymeniderm composed of 19–31 x 8.5–17.5 μm, clavate to subvesiculose, thin- to slightly thick-walled, hyaline to pale yellowish, smooth cells. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 6.0 μm wide hyphae, with pale yellowish brownish walls in KOH. Caulocystidia absent; scattered adpressed to erect, cylindrical to clavate, thin-walled, obtuse terminal cells present (see notes below). Clamp-connections present in all tissues. – Fig. 40

 

Chemical reactions. Hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, on decaying bamboo and Arundinaria leaves, in both montane (alt. 2130–2438 m) and lowland stands.

 

Distribution. Known from Kenya (Pegler 1977), the type locality in the Democratic Republic of Congo, and probably also from Cameroon and Zimbabwe.

 

Revised specimens from tropical Africa.

Cameroon. ? Southwest Province, Korup Forest Reserve, Mundema, 26 March 1991, R. Watling (E).

Democratic Republic of Congo. Lacs Édouard et Kivu District, Albert National Park, Kalonge s. Butahu, 8 May 1953, de Witte 8969 (BR 11375–26, holotype); Ibid., de Witte 8974 (BR 11376–27, isotype).

Kenya. Nyanga province, Kericho District, Kericho, Sambret Tea Estate, W. Mau Forest, 27 March 1968, D.N. Pegler (K(M) 108876).

Zimbabwe. Midland Province, ? Mvuma, Beacon Hill Homestead, 1930 A4, 8 Febr. 1997, C. Sharp 652/97 (BR, n.s.).

 

Notes. Marasmius albertianus is characterised by having a rather small, umbilicate, only slightly sulcate and ochraceous pileus with a darker umbilicus and marginal zone, small ellipsoid basidiospores, clavate or subfusoid marginal cells and by the absence of caulocystidia (but see notes below).

Fungi collected in Kenya by Pegler (1977) have a white pileus with a yellowish brown tinge at centre, a larger stipe (40–110 x 2–4 mm), slightly differently shaped basidiospores (8.5–11.5(–13) x 3.7–5.2 μm, pip-shaped, lacrimoid), shorter cheilocystidia (10–20 x 6.0–10 μm) and well-developed caulocystidia. In my opinion, the form of basidiospores as well as the presence of caulocystidia may be a result of the variability of the species (even in the type specimen, scattered “terminal cells” were found on the stipe surface). The different pileus colour may be caused by the fact that Singer (1964) described this species according to herbarium material, or the collector´s notes (if made!) were too brief. Except for crowded lamellae, Pegler´s description agrees well with a collection by A. Retnowati from Java, Indonesia identified as M. aff. albertianus (Desjardin & al. 2000).

Singer (1964) mentioned in the original description smaller basidiospores (about 7 x 3.2 μm) and the absence of cheilocystidia in comparison with our observations.

Among other African Marasmius species with small basidiospores, M. goossensiae Beeli differs by a larger pileus (25–70 mm), subdistant lamellae, a more robust stipe (3–8 mm wide), smaller basidiospores (6.2–7.9(–9.2) x 3.5–4.2 μm), and by the presence of pleurocystidia; M. witteanus Singer has an umbonate pileus, a densely cespitose-fasciculate stipe, smaller basidiospores ((4.6–)5.0–6.2(–6.6) x (2.5–)3.1–3.8(–4.2) μm), larger cheilocystidia (34.5–85(–135) x (7.0–)10–24(–34.5) μm) and possesses pleurocystidia. In comparison with other species with small basidiospores and without pleurocystidia, M. rhysophyllus Mont. differs by yellow coloured carpophores, intervenose lamellae and smaller basidiospores (5–6.5 x 3.5–4.5 μm, but 6–7 x 4–4.5 μm according to Dennis 1970); M. cohortalis Berk. differs by its pileus being ochraceous at centre and whitish towards margin, intervenose lamellae and smaller basidiospores (6.5–8.0 x 3–4.5 μm, but 5–7 x 2–3 μm according to Dennis 1970) (Pegler 1983; Singer 1976), and M. pellucidus Berk. & Broome by a non-umbilicate, distinctly sulcate, milky white pileus, distant lamellae, a smaller, pruinose to fibrillose stipe (20–55 x 1.5–2 mm), smaller basidiospores (6–7.5 x 2.5–3.5 μm, however, 8–10.5 x 3–4 μm according to Pegler 1986) and developed caulocystidia (Desjardin & Horak 1997).

Collections from Cameroon and Zimbabwe are included with a question mark here because of the absence of a macroscopic description.

 

37. Marasmius kigwenensis Antonín

Antonín, Mycotaxon 85: 116 (2003). Type: Burundi, Bururi Province, Kigwena, Kigwena Forest, 22 Feb. 1979, J. Rammeloo 6709 (BR 11945–14, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 116–118 (2003).

 

Pileus 26–30 mm broad, ± applanate, expanded, with slightly lobate to undulate margin, very thin (± 1 mm), glabrous, ± smooth, somewhat greasy to touch, slightly hygrophanous, pale cream with slightly darker centre, pallescent on drying. Lamellae very crowded, L = ca. 40, l = 2–3, adnexed, narrow (mostly < 1 mm), not intervenose, pale cream (like the pileus), with concolorous entire edge. Stipe 50–70 x 2–3 mm, cylindrical, sometimes slightly laterally compressed, usually somewhat twisted, hollow, densely finely floccose (lens), hard and tough, dark brown (± 7E7 to 7F7) towards base. Context tough, pale watery cream (like pileipellis), with fungoid taste and pleasant, somewhat perfumed smell— Pl. 4

 

Basidiospores 5.8–6.6 x 3.1–3.7 μm, E = 1.6–2.0, Q = 1.8, ellipsoid to lacrimoid, thin-walled, hyaline. Basidia 18.5–20 x 5.8–7.3 μm, 4-spored, clavate. Basidioles 10.0–23 x 3.2–6.9 μm, clavate, cylindrical or subfusoid. Cheilocystidia 15.5–23 x 3.5–5.4 μm, subcylindrical, lageniform, sometimes rostrate or subcapitate, often irregular, rarely branched. Pleurocystidia absent. Hyphae cylindrical to inflated, thin- to slightly thick-walled, up to 20 μm wide. Pileipellis a hymeniderm composed of 17.5–34.5 x 10.0–17.5 μm, clavate to (sub)vesiculose, smooth, ± thin-walled, hyaline cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 9.0 μm wide hyphae, smooth or sometimes with scattered lateral projections, with olivaceous greenish walls in KOH. Caulocystidia 11.5–23 x 6.2–7.9 μm, adpressed to erect, clavate, vesiculose or subcylindrical, slightly thick-walled, greenish in KOH. Clamp-connections present in all tissues.  Fig. 41

 

Chemical reactions. Hyphae and caulocystidia dextrinoid, other structures non-dextrinoid.

 

Ecology. In large groups, single or in small clusters, on rotten wood in a humus layer.

 

Distribution. Known only from the type locality in Burundi.

 

Revised specimens from tropical Africa.

Burundi. Bururi Province, Kigwena, Kigwena Forest, 22 Feb. 1979, J. Rammeloo 6709  (BR 11945–14, holotype).

 

Notes. Marasmius kigwenensis is characterised by a pale coloured pileus, very close lamellae, a finely floccose stipe, small basidiospores, irregular narrow cheilocystidia, the absence of pleurocystidia, the presence of caulocystidia and stipitipellis hyphae turning greenish in KOH.

Among other African Marasmius species with small basidiospores, M. goossensiae Beeli differs by a larger pileus (25–70 mm broad), subdistant lamellae, a more robust stipe (3–8 mm wide), larger basidiospores (6.2–7.9(–9.2) x 3.5–4.2 μm), and by the presence of pleurocystidia; M. witteanus Singer has an umbonate pileus, a densely cespitose-fasciculate stipe, larger cheilocystidia (34.5–85(–135) x (7.0–)10–24(–34.5) μm) and well-developed pleurocystidia; M. arborescens (Henn.) Beeli has densely cespitose stipes, larger basidiospores and larger cheilocystidia; M. albertianus Singer has larger basidiospores, larger cheilocystidia, and no caulocystidia. In comparison with other species with small basidiospores and without pleurocystidia, M. rhysophyllus Mont. differs by yellow coloured carpophores, intervenose lamellae and broader basidiospores (3.5–4.5 μm); M. cohortalis Berk. has its pileus ochraceous at centre and whitish towards margin, intervenose lamellae and larger basidiospores (6.5–8.0 x 3–4.5 μm, but 5–7 x 2–3 μm according to Dennis 1970) (Pegler 1983; Singer 1976), and M. pellucidus Berk. & Broome has a distinctly sulcate, milky white pileus, distant lamellae, a smaller stipe (20–55 x 1.5–2 mm), and larger basidiospores (6–7.5 x 2.5–3.5 μm, however, 8–10.5 x 3–4 μm according to Pegler 1986). Also the white coloured Marasmius nivicola Har. Takah., from Japan, differs especially in larger basidiospores (6–8 x 3–4 μm), larger (9–20 x 5–8 μm), differently shaped cheilocystidia and larger (25–65 x 8–15 μm) caulocystidia (Takahashi 2000a).

  

38. Marasmius favoloides Henn.

Hennings, Bot. Jahrb. Syst. 22: 99 (1895). Type: Cameroon, Yaoundé, Oct. 1894, G. Zenker in: G. Zenker, Flora von Kamerun, No. 468; not preserved (Pegler 1977).

 

Selected descriptions and icons. Hennings, Bot. Jahrb. Syst. 22: 99 (1895); Pegler, Persoonia 4(2): 115 (1966); Pegler, Kew Bull. Addit. Ser. 6: 167–169 (1977); Zoberi, Tropical Macrofungi: 76 (1972).

 

Pileus 15–30 mm broad, at first convex umbonate, soon plane or slightly umbilicate, very thin, lilac grey to cinereous, sometimes light cinnamon-drab at the disc, smooth, strongly radiately ridged, margin entire, undulate. Lamellae adnate to decurrent, cream or with a very pale brownish tint, straight to arcuate, distant but strongly connected by prominent interveining to give a reticulate appearance; edge serrulate. Stipe 20–70 x 1–3.5 mm, equal to attenuated above, hollow, smooth, white pruinose towards the apex, darkening to cinnamon brown below, arising from a white mycelium. Context thin, concolorous. Spore print pure white. (According to Pegler 1977). — Pl. 4

 

Basidiospores (5.2–)6.0–7.0(–7.5) x 3.5–4.5 μm, E = 1.4–1.9, Q = 1.6, ellipsoid, hyaline, smooth, thin-walled; slightly thick-walled crassospores present. Basidia e.g. 30 x 8.0 μm, 4-, rarely 2-spored, clavate; thick-walled crassobasidia present. Basidioles 10–30 x 3.0–8.0 μm, cylindrical or clavate. Cheilocystidia numerous, 10–30 x 7.5–15 μm, (broadly) clavate, subfusoid, pyriform, sometimes irregular, lobed or with 1(–2) projection(s), thin-walled, hyaline. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, hyaline, thin-walled, up to 15 μm wide. Pileipellis a hymeniderm composed of 12–32 x 10–20 μm, vesiculose, (broadly) clavate, subcylindrical, pyriform, sometimes capitate and pedicellate, thin- to slightly thick-walled, smooth cells, with subhyaline to pale ochraceous-yellowish walls in KOH. Pileocystidia absent. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 7.0 μm wide hyphae, with ochraceous-yellow walls in KOH. Caulocystidia 20–48 x 10–22 μm, vesiculose, (broadly) clavate, subfusoid, pyriform, subutriform, thin- to slightly thick-walled, hyaline. Clamp-connections present in all tissues. – Fig. 42

 

Chemical reactions. Basidiospores (except for crassospores) non-dextrinoid, all hyphae, crassospores and crassobasidia dextrinoid.

 

Ecology. Growing on dead leaves and twigs amongst damp forest litter.

 

Distribution. Originally collected in Cameroon (Hennings 1895). Found also in Nigeria, Uganda.

 

Revised specimens from tropical Africa.

Ghana. Tafo, 29 May 1957, M. Holder GC 159 (K(M) 134408).

Nigeria. Ibadan, Ife Biological Gardens, 4 June 1968, M.H. Zoberi 364 (K(M) 134409); Ibid., 12 May 1968, M.H. Zoberi 339 (K(M) 134410).

Uganda. Makerere College, 16 April 1964, E.A. Calder 49 (K(M) 115031).

 

Notes. Marasmius favoloides is characterised by having intervenose to almost reticulate lamellae, rather small basidiospores and well-developed cheilo- and caulocystidia; on the contrary, it lacks pileo- and pleurocystidia.

Among other species with strongly interveined to almost reticulate lamellae, M. rhyssophyllus Mont., found in the Lesser Antilles, has its pileus and stipe in various tinges of yellow, larger cheilocystidia (30–40 x 10–14 μm) and narrower caulocystidia (25–40 x 7–13 μm) (Pegler 1983; Singer 1976); M. cohortalis Berk., from the Lesser Antilles and South America, has a centrally ochraceous brown or yellowish cinnamon and marginally almost whitish coloured pileus, larger cheilocystidia (25–32 x 14–22 μm) and narrower caulocystidia (16–30 x 13–15 μm) (Pegler 1983; Singer 1976). Marasmius poromycenoides Singer, from Argentina and Peru, differs by a smaller (11–16 mm), deep purple pileus, a stipe of the same colour and smaller basidiospores (5.7–6.2 x 3.5–3.8 μm) (Singer 1976).

Although the type specimen is not preserved, Pegler (1966, 1977) is convinced, that his published record really represents this species. In both his descriptions (Pegler 1966, 1977) he described hyphae as “inamyloid though strongly dextrinoid“. However, in notes he mentioned the trama hyphae being only inamyloid and that this species is therefore belonging to sect. Alliacei (= Chordales). The studies of the herbarium specimens from the herbarium Kew showed, that most hyphae are really dextrinoid. Therefore, this species belongs to sect. Globulares. Pegler (1977) in his description mentioned, besides dextrinoid hyphae, the absence of cheilocystidia and smaller basidiospores. 

39. Marasmius violaceoides Antonín

Antonín, Czech Mycol. 56: 249 (2004). Type: Democratic Republic of Congo, Equateur Province, Binga, April 1928, M. Goossens–Fontana 680 (BR 11520–74, as M. violaceus, holotype).

 

Misapplication. Marasmius violaceus Henn. s. Singer 1964, 1965 (= Gymnopus sp.).

 

Selected descriptions and icons. Antonín, Czech Mycol. 56: 249–251 (2004); Singer, Bull. Jard. Bot. Etat Brux. 34: 346 (1964) (as M. violaceus); Singer, Flore Icon. Champ. Congo 14: 263 & Pl. 44, fig. 5. (1965) (as M. violaceus).

 

Pileus 21–40 mm broad, up to 30 mm high, campanulate, then rather applanate, often subumbonate, strongly sulcate except for centre (which is often rugose), striae paler violaceous coloured (striped), sometimes entirely finely veined, glabrous, violaceous, violet brown or brown with slightly violaceous tinge (11D–F5). Lamellae close to subdistant, L = 16–20, l = 1–2, rounded-adnate to adnate, rather narrow, white, whitish or pale greyish orange (± 5B3), with concolorous, entire, curved edge. Stipe 110–125 x 2.5–3.5 mm, slightly attenuated towards apex, hollow, smooth, glabrous, distinctly violaceous above, more distinctly brown towards base; basal mycelium tomentose, dirty white or dirty pale ochraceous. Context thin and whitish in pileus, concolorous with surface and fibrillose, subcartilaginous in stipe, smell and taste fungoid. (According to Singer (1964, 1965a) and a photograph). — Pl. 4

 

Basidiospores 15.5–22.3 x 3.5–5.0 μm, E = 3.7–4.9, Q = 4.3, clavate, cylindrical-clavate to lacrymoid, hyaline; thin-walled. Basidia (28–)35.5–49 x 7.0–10.0 μm, 4-spored, clavate. Basidioles 19–46(–51) x 5.1–11.5 μm, clavate, cylindrical or fusoid. Cheilocystidia 14–24 x (4.2–)6.2–9.2 μm, clavate, thin-walled, hyaline. Pleurocystidia absent. Trama hyphae ± cylindrical, thin-walled, branched, up to 11.5 μm. Pileipellis a hymeniderm composed of 18–40 x 11.5–15(–19) μm, clavate to (sub)vesiculose, thin- to slightly thick-walled, smooth cells, sometimes covered by a thin gelatinous layer. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6 μm wide hyphae, with pale yellowish brownish walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 43

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, on decaying leaves in a marshy forest, on dry soil (“terre ferme”) and in a rain forest with Gilbertiodendron dewevrei.

 

Distribution. Known from Cameroon, the Democratic Republic of Congo, Nigeria and Zambia.

 

Revised specimens from tropical Africa.

Cameroon. South West Province, Korup National Park, trail to Rengo Rock, 8 April 1997, P.J. Roberts K 934 (K(M) 91512; BRNM 691108).

Democratic Republic of Congo. Equateur Province, Binga, April 1928, M. Goossens–Fontana 680 (BR 11520–74, holotype, as M. violaceus); Tshopo Province, close to Batiabongena (5 km NNE), 16 April 1984, B. Buyck 1444 (BR 11753–16); Ibid., B. Buyck 1445 (BR 11752–15).

Nigeria. ? Cross River State, Cross River, Ikom River, 1 May 1990, R.A. Nicholson 418 (K(M) 16722, as M. haematocephalus).

Zambia. Chowo Forest, 10 Dec. 1981, J. Rammeloo 7806 (BR 12019–88); Ibid., 10 Dec. 1981, J. Rammeloo 7757 (BR 12007-76); Ibid., 7 Dec. 1981, J. Rammeloo 7713 (BR 11999–68).

 

Notes. Marasmius violaceoides is characterised by having a distinctly campanulate, distinctly sulcate, violaceous pileus, a very long, violaceous tinged stipe, rather large basidiospores (however, only a few found), very long basidia, clavate cheilocystidia, pileipellis cells sometimes covered by a gelatinous layer, and by the absence of caulocystidia.

From the other violaceous species without developed pleurocystidia, M. musisporus Desjardin & E. Horak has a smaller (8–25 mm), in sulci yellow coloured pileus, a smaller stipe (50–70 x 1–1.5 mm) and very large basidiospores (30–40 x 4.5–5 μm); it was described from Papua New Guinea. Marasmius purpureostriatus Hongo, found in Japan and Papua New Guinea, has a larger (15–50 mm), purple and white to cream striped pileus, a stipe without any violaceous tinge and larger basidiospores (19–28(–32) x 4–6 μm) (Desjardin & Horak 1997, isotype ZT!); M. poromycenoides Singer (Singer 1976), from South America, has a smaller (11–16 mm) reticulate pileus, strongly anastomosed lamellae, a smaller stipe (35–50 x 1–1.5 mm) and small basidiospores (5.7–6.2 x 3.5–3.8 μm).

Singer (1964, 1965a) published this species as M. violaceus Henn. However, he has not revised the type specimen. The type revision showed that M. violaceus represents a collybioid taxon. Therefore, it was necessary to describe M. violaceus s. Singer (1964, 1965a) as a new species.

                                                                                                                                                       

40. Marasmius mesosporus Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 349 (1964). Type: Democratic Republic of Congo, Equateur Province, Binga, Aug. 1947, M. Goossens–Fontana 4065 (BR 11497–51, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 349–350 (1964); Singer, Flore Icon. Champ. Congo 14: 265 & Pl. 44, fig. 9 (1965).

 

Pileus 15–25 mm broad, campanulate, then campanulate-convex, with a small umbilicus, strongly striate, glabrous, pale violaceous, with ochraceous centre. Lamellae distant to moderately distant, L = 15–16, l = 2, adnate to free, rather broad, slightly intervenose or not, white to very pale lilac. Stipe 140–150 x 2–3 mm, cylindrical, slightly broadened and curved at base, hollow, glabrous, smooth, yellowish brown towards apex, darker brown towards base; basal mycelium tomentose, pale ochraceous, with a transition to membranaceous mycelium covering the substrate. Context subconcolorous and very thin in pileus, concolorous and subcartilaginous in stipe, elastic at base, with acrid taste and smell. (According to Singer (1964, 1965a), original description by Goossens-Fontana). — Pl. 5

 

Basidiospores (according to Singer, not found by me in type specimen) 10.5–13.5(–15.5) x 4.5–5.3(–6.7) μm, fusoid-oblong, hyaline, smooth. Basidia 30–36 x 7.7–10.5 μm, 4-spored, clavate, some of them with refractive granular contents. Basidioles 17–35 x 4.0–10.0 μm, cylindrical, clavate to (sub)fusoid, sometimes subrostrate or subcapitate. Cheilocystidia 22–28 x 12–16 μm, similar to pileipellis cells, mixed with basidiola-like elements. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, up to 15 μm wide. Pileipellis a hymeniderm composed of 19–30 x 11–19 μm, (broadly) clavate, subfusoid to subsphaeropedunculate, smooth, thin- to slightly thick-walled cells with (sub)hyaline walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, thick-walled, smooth, up to 7.0 μm wide hyphae. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 44

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, in humus in a dry forest.

 

Distribution. Known only from the type locality in the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Equateur Province, Binga, Aug. 1947, M. Goossens–Fontana 4065 (BR 11497–51, holotype).

 

Notes. Marasmius mesosporus is characterised by having a rather small, pale violaceous, sulcate pileus, pale lilac lamellae, a very long stipe, an acrid taste and smell, cheilocystidia similar to pileipellis cells and by the absence of pleuro- and caulocystidia. Some of the basidioles have granular refractive contents, however, their size perfectly fits in the size variability of the other, “true” basidioles.

Among other tropical violaceous tinged species, M. musisporus Desjardin & E. Horak has a smaller stipe and distinctly larger basidiospores (30–40 x 4.5–5 μm), and M. purpureostriatus Hongo (ZT 3221, isotype!) has larger carpophores, basidia (45–50 x 11–15 μm) and basidiospores (20–31 x 5.0–6.0 μm).

  

41. Marasmius zenkeri Henn.

Hennings, Bot. Jahrb. Syst. 22: 98 (1895). Type: Cameroon, Yaoundé Station, 2 April 1894, Zenker & Staudt 286 (BR 18910–92, ex B, lectotype). – Marasmius superbus Henn., Bot. Jahrb. Syst. 30: 48 (1901).

       

Selected descriptions and icons. Hennings, Bot. Jahrb. Syst. 22: 98 (1895); Pegler, Kew Bull. Addit. Ser. 6: 171–173 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 348–349 (1964); Singer, Flore Icon. Champ. Congo 14: 264–265 & Pl. 44, fig. 3 (1965); Zoberi, Tropical Macrofungi; 70–72. (1972).

 

Pileus 30–100 mm broad, convex, then convex-applanate, slightly umbonate or subumbonate, deeply sulcate, glabrous, smooth or slightly rugulose at centre, lilac pinkish, dark lilac-violaceous (always at centre), often rather dirty pinkish, pale violaceous or pale on sulci and then radially striped, without yellow tinge. Lamellae distant or very distant, L = 13–19, l = 3, adnexed or free, slightly ventricose, narrow or moderately broad (up to 10 mm), not intervenose, pale lilac, with paler or pale edge. Stipe (40–)120–170 x 2–12 mm, cylindrical, with broadened base, or attenuated towards apex (to 2.5–5 mm), often twisted, hollow, smooth, glabrous, brown then fuligineous, dark reddish brown or chestnut brown towards base, often entirely fuligineous in the end; basal mycelium woolly, silky-tomentose, strigose, pale ochraceous, dirty greyish or whitish. Context white, tough in pileus, pale lilac under pileipellis in centre, pale to almost concolorous under stipitipellis, without smell, with a mild, then slightly acrid after-taste. Spore print white or pale lilac. (According to Pegler (1977) and Singer (1964, 1965a)).

 

Basidiospores (15.5–)17.723(–27) x 4.2–5.6 μm, E = 3.2–5.3, Q = 3.74.4, lacrymoid, narrowly fusoid to narrowly clavate, sometimes curved, smooth, thin-walled, hyaline. Basidia 30–39 x 7.5–12 μm, 4-spored, clavate. Basidioles 15.5–42 x 4.0–12 μm, clavate, subcylindrical, subfusoid. Cheilocystidia 14–50 x 10–15 μm, clavate, thin- to slightly thick-walled, hyaline. Pleurocystidia absent. Trama hyphae cylindrical, thin-walled (slighthly thick-walled in subpileipellis), up to 20 μm wide. Pileipellis a hymeniderm composed of (16–)19–32(–40) x 10–19(–23) μm, clavate, mostly slightly thick-walled (at least in upper part), smooth cells, with brownish walls in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 8 μm wide hyphae. Caulocystidia absent; scattered, adpressed to erect, cylindrical terminal cells present on stipe surface. Clamp-connections present in all tissues. – Fig. 45

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid;

 

Ecology. Single or in pairs on litter (e.g. of Brachystegia) in a forest on dry soil (terre ferme) and in a montane rainforest.

 

Distribution. Known from the type locality in Cameroon, the Democratic Republic of Congo, Ivory Coast, Malawi, Uganda and probably also Tanzania.

 

Revised specimens from tropical Africa.

Cameroon. Yaoundé Station, 2 April 1894 (1890–94), G. Zenker & Staudt 286 (BR 18910–92, lectotype; K(M) 108849, K (s.n.), PC (s.n.), PRM 707416 ex B, syntypes); Bipinde, Urwaldgebiet, 1900, G. Zenker in: Zenker, Flora von Kamerun, No. 2214 (K(M) 134629, PC, as M. zenkeri f. cinerascens); Ibid., 1899, G. Zenker, in: Zenker, Flora von Kamerun, No. 2021 (K(M) 134630); Ibid., Aug. 1899, G. Zenker 2172 (S F–16257, syntype of M. superbus).

Democratic Republic of Congo. Equateur Province, Binga, Sept. 1927, M. Goossens–Fontana 626 (BR 11534–88); Ibid., Oct. 1944, M. Goossens–Fontana 3046 (BR 11536–90); Ibid., Sept. 1936, M. Goossens–Fontana 1046 (BR 11535–89).

Ivory Coast. Abidjan, Forêt du Banco, 22 May 1976, L. Aké Assi 480 (K(M) 134633).

Malawi. Makwawa, Zomba, 10 Jan. 1980, B. Morris 56 (K(M) 134634).

Tanzania. ? West Usambara Mts., Shagayu Forest Reserve, 5 km NW of Mlalo Mission, 15 Febr. 1985, I. Krisai (WU).

Uganda. Buganda Province, Mengo District, Mawacota County, Mpanga Research Forest, 7 June 1968, D.N. Pegler U 1234 (K(M) 134631).

 

Notes. Marasmius zenkeri is characterised by having very large and long-stiped carpophores with a lilac, often radially striped, deeply sulcate pileus, distant, pale lilac lamellae, a fuligineous stipe, clavate cheilocystidia, and by the absence of pleurocystidia and caulocystidia. The size of the cheilocystidia is very variable: I measured 14–21 x 7.7–11.5 μm, 15–55 x 10.0–15.5 μm, and 34.5–50 x 11.5–19 μm cheilocystidia in various collections.

Singer (1964, 1965a) mentioned narrower basidiospores (x 3.5–4.0 μm), and smaller pileipellis cells (e.g. 18 x 11 μm), Pegler (1977) measured smaller cheilocystidia (15–18 x 5–7 μm) and smaller pileipellis cells (18–20 x 5–7 μm). According to Hennings (1895) it represents an edible fungus.

All other violaceous species without pleurocystidia especially differ in having smaller carpophores (pileus smaller than 25 mm) and distinctly different basidiospores. Only M. purpureostriatus Hongo, described from Japan and also known from Papua New Guinea, has an up to 50 mm broad pileus. It differs by the different pileus colour (purple at centre and on sulci, white to cream elsewhere), cream coloured lamellae, a fibrillose stipe surface, slightly larger basidiospores (19–28(–32) x 4–6 μm) and slightly smaller cheilocystidia (15–37 x 10.0–15 μm) (Desjardin & Horak 1997, isotype ZT!).

  

42. Marasmius bekolacongoli Beeli

Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928). Type: Democratic Republic of Congo, Equateur Province, Eala, Oct. 1923, M. Goossens–Fontana 204 (BR 11406–57, holotype).

 

Selected descriptions and icons. Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928); Pegler, Kew Bull. Addit. Ser. 6: 171–172 (1977); Ryvarden, Piearce & Masuka, Larger Fungi of South Central Africa: 96–97 (1994); Singer, Bull. Jard. Bot. Etat Brux. 34: 346–347 (1964); Singer, Flore Icon. Champ. Congo 14: 263–264 & Pl. 44, fig. 6 (1965).

 

Pileus (10–)30–67(–100) mm broad, campanulate, then applanate-convex, with depressed centre, strongly sulcate, translucently striate, glabrous, never rugulose, crenulate at margin, violaceous brown (11E5-11F5–6) when young, then dull red or greyish red (11C3–11E4) at centre and on striae, yellowish white, greyish yellow (4A2–AB3) to lemon yellow elsewhere. Lamellae distant to very distant, L = 7–20, adnate to almost free, rather broad, not intervenose but often rugulose between lamellae, yellowish white or pale lemon yellow (2A2–3), with whitish edge. Stipe 50–150 x 2.5–6(–10) mm, cylindrical or slightly attenuated towards apex, hollow, glabrous, smooth, longitudinally grooved when old, pale yellow when young (4A3), then pale yellow to greyish yellow (4A3–4B3) at apex, (sometimes?) with a lilac tinge when young, then dark blond to light brown (5D4–5) and brown (± 7D6); basal mycelium white, tomentose, transient to a membranaceous layer on the substrate. Context very thin, hollow, pale lilac to brown, with fungoid smell, with strong and mild to bitter taste. Spore print white. (According to Pegler (1977), Singer (1964, 1965a), some additions according to dry specimens). — Pl. 6

 

Basidiospores 17.5–24.5(–26) x (3.8–)4.2–5.4 μm, E = 4.2–6.1, Q = 5.1, clavate, subcylindrical, often slightly curved, thin-walled, hyaline. Basidia 30–38.5 x 8.0–10.0 μm, 4-spored, clavate. Basidioles 15.5–38.5 x 3.8–10.0 μm, clavate to subcylindrical. Cheilocystidia 15.5–24 x (6.9–)8.5–14 μm, clavate to subfusoid, thin- to slightly thick-walled. Pleurocystidia absent. Hyphae ± cylindrical, thin-walled, slightly thick-walled in subpileipellis, up to 12 μm wide. Pileipellis a hymeniderm composed of 15.5–27(–31) x (7.0–)11.5–19 μm, clavate, broadly clavate to vesiculose, thin- to slightly thick-walled, smooth cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae, subhyaline to brownish in KOH. Caulocystidia absent; scattered lateral projections or adpressed to suberect terminal cells present on stipe surface. Clamp-connections present in all tissues. – Fig. 46

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid (pileipellis cells sometimes seem to be slightly dextrinoid).

 

Ecology. Single, on litter in an inundated forest, a Miombo woodland with Brachystegia microphylla, B. glaucescens and B. utilis, on leaves of Canarium schweinfurthii, an evergreen forest with Combretum and litter of an oil-palm.

 

Distribution. Known from Burundi, Cameroon, the Democratic Republic of Congo, Kenya, Malawi, Nigeria, Tanzania, Uganda and Zimbabwe. Outside of tropical Africa, it it mentioned from China (Li Taihui & al. 1990), but this collection may represent a misidentification (see notes below).

 

Revised specimens from tropical Africa.

Burundi. Rumonge Province, Nkayamba near Rumonge, 15 March 1994, A. Verbeken 94/206 (BR 27280–23); Bururi Province, Kigwena, Kigwena Forest, 22 Feb. 1979, J. Rammeloo 6721 (BR 11950–19).

Cameroon. South West Province, Korup National Park, Ireba–Inene Camp, 14 April 1997, P.J. Roberts K 1176 (K(M) 91524; BRNM 691107); Ibid., trail to Rengo Rock, 1 May 1996, P.J. Roberts 290 & 291 (K(M) 39425); Ekombitié, Mbalmayo Forest Reserve, 26 Apr. 2001, C. Douanla–Meli 058 (HUYI).

Democratic Republic of Congo. Equateur Province, Binga, June 1947, M. Goossens–Fontana 4052 (BR 11407–58); Equateur Province, Eala, Oct. 1923, M. Goossens–Fontana 204 (BR 11406–57, holotype); Kalonga, Katavuleko, 6 May 1953, H.F. de Witte 8941 (BR 11408–59).

Kenya. Nyanza Province, Kericho District, Man Forest, African Highlands Estate, Kiptiget River, Kericho, 26 March 1968, D.N. Pegler K 264 (K(M) 8824); ? Coast Province, Kwale District, Mrima Hill, 10 April 1978, B. Verdcourt 5275A (K(M) 134411); Coast Province, Kwale District, Shimba Hills, Mt. Dzombo, 8 April 1968, F. Magogo & P. Glower 788 (K(M) 134412).

Malawi. Ruo Gorge, Mulanje, 20 Dec. 1981, B. Morris 373 (K(M) 134414); Southern Province, Thyolo District, Naminkweya Estate, 3–4 March 1973, L. Ryvarden 11204 (K(M) 134415).

Nigeria. Cross River State, Calabar–Cameroon Road, 23 June 1990, R.A. Nicholson 535 (K(M) 17067).

Tanzania. Dar es Salaam District, Spring 1970, J.N.R. Kasembe K 307 (K(M) 134413).

Uganda. Buganda Province, Mengo District, Mawakota County, Mpanga Research Forest, 13 June 1968, D.N. Pegler U 1459 (K(M) 8823).

Zimbabwe. Mvuma, Lovedale Ranch, hill north of road in front of “incubator”, 1930A4, 6 Febr. 1999, A. Verbeken 99–135 (GENT, BRNM).

 

Notes. Marasmius bekolacongoli is characterised by having a large, violaceous pileus with lemon yellow stripes, distant yellowish lamellae, a very large stipe with well-developed basal mycelium and a bitter taste. Microscopically, it has large basidiospores, clavate to subfusoid cheilocystidia, and lacks developed caulocystidia. Collections P.J. Roberts 290 & 291 agree well in size and microfeatures, however, its stipe is dark purple (especially at base) and the striation pale violaceous to cream.

A description of microfeatures by Pegler (1977) agrees with ours, a description by Singer (1964, 1965a) differs only by narrower basidiospores (x 3.0–4.5 μm). Li Taihui & al. (1990) probably misidentified this species because he included it in sect. Sicci. Collection Goossens-Fontana 4052 differs microscopically in having broader basidiospores (19.5–22.5 x 5.4–6.4 μm, E = 3.1–4.3, Q = 3.5).

Marasmius purpureostriatus Hongo differs especially in having less robust carpophores (pileus 15–50 mm broad, stipe 70–115 x 1.5–2.5 mm), a white to cream (not lemon yellow) striped pileus, a fibrillose stipe surface, slightly larger basidiospores (19–28(–32) x 4–6 μm); it is known from Japan and Papua New Guinea (Desjardin & Horak 1997, isotype ZT!). Other violaceous coloured species without pleurocystidia are distinctly smaller (pileus up to 25 mm broad) and have distinctly smaller or larger basidiospores.

Beeli (1928a) described M. bekolacongoli var. brunneolus Beeli and mentioned two collections by M. Goossens–Fontana; coll. 115 represents M. brunneolus (Beeli) Singer and coll. 151 M. flavus Singer.

This fungus and M. brunneolus are known under the local name “becolakongoli” (Hendrickx 1948). On the specimen label of the collection by Magogo & Glower from Kenya, following note is added: “said to be poisonous and causing intoxications and vomiting, local name: uwoga koma”.

 

43. Marasmius brunneolus (Beeli) Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 344 (1964). – Marasmius bekolacongoli var. brunneolus Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928). Type: Democratic Republic of Congo, Equateur Province, Eala, May 1923, M. Goossens–Fontana 115 (BR 11414–65, holotype). – Marasmius staudtii var. pallida Henn., Bot. Jahrb. Syst. 22: 97 (1895) (according to Singer).

 

Misapplication. Marasmius brunneolus (Berk. & Broome) Pegler, Kew Bull. Add. Ser. 12: 163 (1986) (= M. davidii Antonín 2003).

 

Selected descriptions and icons. Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928); Singer, Bull. Jard. Bot. Etat Brux. 34: 344–346 (1964); Singer, Flore Icon. Champ. Congo 14: 263 & Pl. 44, fig. 1 (1965).

 

Pileus 30–80 mm broad, convex, umbilicate or not, then convex-applanate, obtuse, deeply sulcate, glabrous, fuligineous brownish, greyish brown to reddish brown (7D3–5), rarely paler, with whitish, yellowish white or yellowish grey (4A2, 4B2) striae. Lamellae subdistant to distant, L = ca. 15–20, l = 2, emarginate, obtuse or attenuate, narrowly adnexed to almost free, rather narrow, with lamellulae, not intervenose, ivory whitish, cream, pale yellow (3A3), pale alutaceous, sometimes with pinkish tinge, with concolorous entire edge. Stipe 70–180 x 3–6 mm, cylindrical, with slightly broadened base and attenuate apex (up to 15 mm), often curved or twisted, hollow, glabrous, longitudinally striate, pale yellow (3A3) above, concolorous with pileus or often darker, brown-red, brown (7D3–5), sepia, dark chestnut brown (8E–F6); basal mycelium strigose or tomentose, dirty cream, fulvous-ochraceous or whitish, transient to a cream to alutaceous mycelium covering decaying leaves as a thin fibrillose layer. Context white, often concolorous with surface in stipe, with acrid smell and taste— Pl. 7

 

Basidiospores 15.5–25.5 x 3.8–5.4 μm, E = 4.0–5.4, Q = 4.4–4.6, clavate, lacrimoid, subfusoid to cylindrical-clavate, hyaline, smooth. Basidia 31–38.5 x 10.0–14.5 μm, 4-spored, clavate. Basidioles 16–45(–48) x 4.0–12 μm, clavate to cylindrical. Cheilocystidia 14–23.5(–27) x 8.0–11.5(–17) μm, clavate, thin-walled, rarely slightly thick-walled, hyaline. Pleurocystidia not found. Hyphae cylindrical to subinflated, thin- to slightly thick-walled, up to 12 μm wide. Pileipellis a hymeniderm composed of 15.5–24.5(–35) x 10.5–16(–19) μm, clavate to subvesiculose, thin- to slightly thick-walled, smooth cells, hyaline to pale yellowish brownish in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 47

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. On litter in a virgin forest, on dry soil (terre ferme), in a marshy forest, and in a primary forest with Gilbertiodendron and Scaphopetalum, but also in ruderalised scrub.

 

Distribution. So far known only from the Democratic Republic of Congo, Zimbabwe and possibly Nigeria.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Equateur Province, Eala, May 1923, M. Goossens–Fontana 115 (BR 11414–65, holotype); Yangambi, Isalowe, 18 May 1939, Louis 14870 (BR 11416–67); Equateur Province, Binga, Aug. 1947, M. Goossens–Fontana 4053 (BR 11415–66); Tshopo Province, Kisangani, NNE to Batiabongena, 27 April 1984, B. Buyck 1566 (BR 11741–04).

Nigeria. Akwa Ibom State, Anua Akira, St. Luke´s Hospital, 14 April 1989, R.A. Nicholson 148 (K(M) 7512, as M. mesosporus).

Zimbabwe. Chipinge, Busi–farm, forests around Scott´s House – 2032 B1, 12 Febr. 1999, A. Verbeken 99–209 (GENT, BRNM).

 

Notes. Marasmius brunneolus is characterised by having very large and long-stiped carpophores, a fuligineous brownish pileus, an acrid taste, large basidiospores, large basidia, clavate cheilocystidia and by the absence of caulocystidia.

Singer (1964, 1965a) mentioned slightly narrower basidiospores (x 2.7–4.5 μm) and smaller pileipellis cells (about 15 x 10 μm).

Marasmius tshopoensis Antonín seems to be the closest related species. It differs by an ochraceous, uniformly coloured pileus, no lamellulae, larger basidiospores ((19–)21–26 x 4.8–6.0 μm), and more variable cheilocystidia. Marasmius riparius Singer, described from South America, has a smaller pileus (25–45 mm) coloured like M. oreades, a shorter stipe (20–60 x 1–3 mm), smaller basidiospores (8.2–10 x 4.8–5.5 μm) and well-developed pleurocystidia (Singer 1976). Marasmius cohortalis Berk., from the Lesser Antilles, has a 10–25 mm broad pileus which is ochraceous brown at centre and whitish towards margin, strongly intervenose lamellae, a short stipe (18–50 x 1–3 mm), small basidiospores (6.5–8 x 3–4.5 μm) and smaller cheilocystidia (25–32 x 14–22 μm); M. plumieri (Lév.) Singer has a rusty brown, 30–70 mm broad pileus, brown or glaucous lamellae, a smaller stipe (40–80 x 2–4 mm) and smaller basidiospores (14.5–15.5 x 4–4.5 μm); both of them grow in the Lesser Antilles (Pegler 1983). Marasmius ochraceus Berk. & Broome, from Sri Lanka, differs by its non-sulcate pileus, small basidiospores (5.5–7.2 x 2.7–3.5 μm), the absence of cheilocystidia and the presence of gloeocystidia; M. calvus Berk. & Broome, also from Sri Lanka, by a smaller pileus (20–45 mm) and stipe (45–80 x 2–4 mm), smaller basidiospores (8–10.5 x 2.7–3.5 μm), smaller basidia (20–22 x 4–5 μm), the absence of cheilocystidia and the presence of gloeocystidia.

Marasmius brunneolus is known under the local name “becolakongoli” (Hendrickx 1948).

  

44. Marasmius tshopoensis Antonín

Antonín, Mycotaxon 85: 128 (2003). Type: Democratic Republic of Congo, Tshopo Province, close to Kisangani, Isle of Congolo, 14 April 1984, B. Buyck 1394 (BR 11762–25).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 128–130 (2003).

 

Pileus 60 mm broad, somewhat applanate, strongly sulcate, glabrous, membranaceous, with straight entire margin, rather uniformly coloured, never striped, yellow ochraceous at centre, ochraceous on sulci, striae paler, almost cream coloured towards margin, ochraceous when dry. Lamellae very distant, L = 17, l = 0, free, strongly intervenose, about 8 mm broad, yellowish white (2A2), with concolorous entire edge. Stipe 85 x 2.5 mm, cylindrical, slightly twisted, glabrous, but locally with arachnoid network of small fibrils, hollow, smooth, dark brown (7F4); with rich white basal tomentum. Context thin, yellowish white (2A2), not changing colour; with slightly sweet taste and smell. Spore print white. — Pl. 7

 

Basidiospores (19–)21–26 x 4.8–6.0 μm, E = 3.8–4.8, Q = 4.3, clavate, narrowly fusoid, rarely septate, thin-walled, hyaline. Basidia ca. 33 x 9.0 μm, 4-spored, clavate. Basidioles 15.5–35.5 x 3.0–10.0 μm, clavate, cylindrical or subfusoid. Cheilocystidia (12.5–)15.5–28.8 x 6.2–11.5(–21) μm, variable in shape, clavate, subvesiculose, cylindrical, sometimes irregular, lobed or subrostrate, rarely subcoralloid, ± thin-walled, hyaline. Pleurocystidia absent. Hyphae ± cylindrical, thin- to slightly thick-walled (in subpileipellis), up to 11 μm wide. Pileipellis a hymeniderm composed of 15.5–25.5 x 10–19(–24.5) μm, clavate to vesiculose, thin- to slightly thick-walled, smooth cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 6.0 μm wide hyphae. Caulocystidia absent; scattered adpressed to erect terminal cells present. Clamp-connections present in all tissues. – Fig. 48

 

Chemical reactions. All hyphae dextrinoid, pileipellis cells sometimes slightly dextrinoid, other structures non-dextrinoid.

 

Ecology. On fallen decaying leaves on humus in a secondary rain forest with relicts of a primary rain forest.

 

Distribution. So far known only from the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Tshopo Province, close to Kisangani, Isle of Congolo, 14 April 1984, B. Buyck 1394 (BR 11762–25).

 

Notes. Marasmius tshopoensis is characterised by having rather large carpophores, a sulcate pileus with yellow-ochraceous centre, ochraceous sulci and almost cream marginal area, distant, distinctly intervenose pale yellowish lamellae, absent lamellulae, large basidiospores, rather small cheilocystidia, and by the absence of pleuro- and caulocystidia.

Marasmius brunneolus (Beeli) Singer is the closest related species. It differs by an often whitish sulci, present lamellulae, smaller basidiospores (15.5–25.5 x 3.8–5.4 μm), and more uniformly clavate cheilocystidia. For a detailed discussion see notes on M. brunneolus.

Among other species with strongly intervenose lamellae, Marasmius rhyssophyllus Mont. has a smaller (25 mm) yellow pileus, yellow lamellae and a yellow or white stipe, and smaller basidiospores, 7.0–8.3 x 3.3–3.8 μm (5.0–6.5 x 3.5–4.5 μm, according to Pegler 1983); M. ditopotrama Singer has a light brownish grey pileus with otter brown or dark spadiceous striae, brownish or grey lamellae, a pruinose, grey stipe, smaller basidiospores (7–9 x 4.5–4.8 μm) and larger cheilocystidia; M. poromycenoides Singer has a dark purple pileus and stipe and smaller basidiospores (5.7–6.2 x 3.5–3.8 μm), and M. cohortalis Berk. (more varieties are described) has smaller basidiospores (up to 8.5 μm long) in all varieties. All above mentioned species are known from South America (Dennis 1970; Singer 1976), the first and also the last one from Sri Lanka.

  

45. Marasmius missangoënsis Pat.

Patouillard, Bull. Soc. Mycol. Fr. 18: 299 (1902). Type: Democratic Republic of Congo, village of Missango, Bords de l´Oubangui, 1891, Dybowski 488 (K(M) 92589, isotype ?). – Marasmius schweinfurthianus Henn., Bot. Jahrb. Syst. 17: 32 (1893).

 

Selected descriptions and icons. Patouillard, Bull. Soc. Mycol. Fr. 18: 299 (1902); Saccardo & D. Saccardo, Syll. fung. 17: 42–43 (1905).

 

Pileus 20–100 mm, hemispherical at first, then applanate, thin, membranaceous, glabrous, with sinuate margin, distinctly and deeply radially sulcate, rusty coloured (rufus). Lamellae slightly ventricose, broad, attenuate, adnate, intervenose. Stipe 120 x 5–6 mm, terete, coriaceous, longitudinally striate, pruinose, attenuate towards apex, brown-black. (According to Patouillard 1902).

 

Basidiospores 23.5–28 x 5.5–7.0 μm, E = 3.7–5.4, Q = 4.2, clavate, narrowly fusoid, sublacrimoid, thin-walled, hyaline. Basidia not found. Basidioles up to 45 x 9.0 μm, cylindrical, clavate, fusoid. Cheilocystidia 15–18 x 9.0–12 μm, (broadly) clavate, thin-walled. Pleurocystidia absent. Trama hyphae cylindrical, thin-walled, branched, up to 12 μm wide. Pileipellis a hymeniderm composed of 18–25 x 11–15 μm, clavate, thin- to slightly thick-walled, smooth cells. Stipe absent in type specimen. Clamp-connections present in all tissues. – Fig. 49

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. On soil.

 

Distribution. Known only from the Democratic Republic of Congo and probably also Tanzania.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Village of Missango, Bords de l´Oubangui, 1891, Dybowski 488 (K(M) 92589, isotype ?).

Tanzania. (as Central Africa) ? Seengebiet, Wakondjo, 1 Jan. 1892, G. Schweinfurth (S F6260, syntype of M. schweinfurthianus).

 

Notes. Marasmius missangoënsis is characterised by having a rusty coloured pileus, large basidiospores, long basidioles, small cheilocystidia and by the absence of pleurocystidia.

Marasmius caryotae (Berk.) Petch differs especially by a straw yellow to greyish yellow pileus, smaller basidiospores (19–25 x 5–6.2 μm) (Pegler 1986); M. viridis Desjardin & E. Horak differs by its green coloured and distinctly smaller (10–15 mm) pileus, paler stipe and narrower basidiospores (20–25 x 4–5 μm); M. amabilis Hariot & Pat. in having a smaller (20–25 mm), white to cream buff pileus; M. purpureostriatus Hongo has a purple pileus disc and furrows and narrower basidiospores (19–28 x 4–6 μm) (Desjardin & Horak 1997).

 

46. Marasmius witteanus Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 354 (1964). Type: Democratic Republic of Congo, Kalonga, Katavuleko, 6 May 1953, de Witte 8934 (BR 11521–75, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 354. 1964; Singer, Flore Icon. Champ. Congo 14: 266–267. 1965.

 

Pileus 8–18 mm broad, convex, umbonate, applanate to slightly depressed around umbo, glabrous, smooth, ochraceous brown when dry. Lamellae close, L = ca. 30–35, l = 3, free, rather narrow, pale brown when dry. Stipe 50–110 x 1–2 mm, cylindrical, densely cespitose-fasciculate, subglabrous, fuligineous; basal mycelium tomentose, penetrating substrate. Context thin at margin. (According to Singer (1964, 1965a)).

 

Basidiospores (4.6–)5.0–6.2(–6.6) x (2.5–)3.1–3.8(–4.2) μm, E = 1.3–1.8(–2.0), Q = 1.6–1.8, ellipsoid to broadly ellipsoid, both thin-walled non-dextrinoid and slightly thick-walled (crassospores) dextrinoid, smooth. Basidia 15.5–28 x 4.6–7.3 μm, 4-spored, clavate; crassobasidia present. Basidioles 11.5–29(–31) x 3.3–9.0(–11.5) μm, clavate, cylindrical. Cheilocystidia 34.5–85(–135) x (7.0–)10–24(–34.5) μm, variable, often voluminous, cylindrical, clavate, fusoid, often subrostrate, hyaline, with refractive contents, thin-walled. Pleurocystidia similar to cheilocystidia. Hyphae cylindrical to subinflated, thin- to slightly thick-walled (often in subpileipellis), pale yellowish brownish, up to 19 μm wide. Pileipellis a hymeniderm (sometimes loosening) composed of 14–24(–32) x (6.2–)8.5–14(–19) μm, clavate, thin- to slightly thick-walled, smooth cells yellowish, yellowish brownish or brownish in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 7 μm wide hyphae. Caulocystidia absent. Clamp-connections present in all tissues.   Fig. 50

 

Chemical reactions. All hyphae and crassospores dextrinoid, other structures non-dextrinoid.

 

Ecology. Densely cespitose, on decaying leaves and wood conglomerated together by mycelium in wet places in a shaded montane forest.

 

Distribution. Known only from the type locality in the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Kalonga, Katavuleko, 6 May 1953, de Witte 8934 (BR 11521–75, holotype); Ibid., de Witte 8943 (BR 11529–83); Ibid., de Witte 8935 (BR 11526–80); Ibid., de Witte 8937 (BR 11524–78); Ibid., de Witte 8936 (BR 11525–79); Ibid., de Witte 8938 (BR 11523–77); Ibid., de Witte 8939 (BR 11527–81); Ibid., de Witte 8940 (BR 11522–76); Ibid., de Witte 8944 (BR 11528–82); all isotypes.

 

Notes. Marasmius witteanus is characterised by having a very small, non-striate pileus on a long, densely cespitose stipe, very small, ellipsoid to broadly ellipsoid, both thin-walled non-dextrinoid and slightly thick-walled dextrinoid basidiospores, variable, often very large and voluminous pleuro- and cheilocystidia, and by the absence of caulocystidia. This combination of features is rather unique in the genus Marasmius.

Among tropical species with small basidiospores (less than 10 μm long) and well-developed pleurocystidia, M. goossensiae Beeli differs especially in having larger and paler coloured carpophores, a thicker stipe (3–6(–8) mm), and larger basidiospores (6.2–7.9(–9.2) x 3.5–4.2 μm). Marasmius ochraceus Berk. & Broome has a large pileus (30–80 mm) and a thicker stipe (60–110 x 4–7 mm); M. calvus Berk. & Broome a campanulate pileus, larger differently shaped basidiospores (8–10.5 x 2.7–3.5 μm, cylindrical to subfusoid) and smaller refractive cystidia (30–35 x 6–8 μm). Both species are known from Sri Lanka (Pegler 1986). Marasmius phlebodiscus Desjardin & E. Horak, from Papua New Guinea, has a pale beige to tan coloured, 30–60 mm large pileus with net-like wrinkles at centre, and only fusoid pleurocystidia (Desjardin & Horak 1997). Marasmius riparius Singer, from Argentina, has a differently coloured pileus (like M. oreades), a white then cinnamomeous stipe, larger (8.2–10 x 4.8–5.5 μm) ellipsoid-fusoid basidiospores and smaller cystidia (40–44 x 8–9 μm); M. silvicola Singer has a larger (17–95 mm) deeply cinnamomeous to ochraceous brownish pileus, a larger stipe (20–118 x 1.5–7 mm), and cheilocystidia different from pleurocystidia (Singer 1976).

A description by Singer (1964, 1965a) differs only in slightly smaller basidia (23–24 x 5.2–5.5 μm) and hymenial cystidia (22–63 x 9.0–20 μm).

 

47. Marasmius goossensiae Beeli

Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928). Type: Democratic Republic of Congo, Equateur Province, Binga, Nov. 1925, M. Goossens–Fontana 397 (BR 11453–07, lectotype).

 

Selected descriptions and icons. Beeli, Bull. Soc. Roy. Bot. Belg. 60: 157 (1928); Pegler, Kew Bull. Addit. Ser. 6: 176 & 178 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 352–353. (1964); Singer, Flore Icon. Champ. Congo 14: 266 & Pl. 45, fig. 4 (1965); Zoberi, Tropical macrofungi: 80–81 (1972).

 

Pileus 25–70 mm broad, convex, then with applanate to revolute margin, ± applanate to convex-subumbonate, but never really umbilicate or umbonate in the end, later distinctly striate at margin, glabrous, cream, with fuligineous brown or ochraceous brown centre, and avellaneous grey, grey cream or cream striae. Lamellae subdistant, emarginate to almost free, often finely intervenose, moderately broad (6–10 mm near stipe), subconcolorous with pileus margin. Stipe 50–70(–100) x 3–6(–8) mm, cylindrical, but often slightly broadened or attenuated at base, hollow, (sub)glabrous, smooth (however, it seems to be longitudinally striate when dry), concolorous with pileus, often more cream above and more fuligineous at base; basal mycelium present. Context whitish and thin at margin in pileus, concolorous with surface in stipe. Spore print white. (According to Beeli (1928a), Singer (1964, 1965a)). — Pl. 8

 

Basidiospores 6.2–7.9(–9.2) x 3.5–4.2 μm, E = 1.6–2.2, Q = 1.8, ellipsoid, thin-, rarely slightly thick-walled, smooth, hyaline. Basidia 23–26 x 6.5–8.5 μm, 4-spored, clavate. Basidioles 15.5–31 x 4.5–8.5 μm, clavate to cylindrical. Cheilocystidia 24–31(–38.5) x 10–14 μm, clavate to subfusoid, thin-walled, hyaline in KOH. Pleurocystidia 48–100 x 12.5–20(–23) μm, clavate, fusoid, often rostrate, thin- to slightly thick-walled, (sub)hyaline. Hyphae cylindrical to subinflated, sometimes consisting of ± short ellipsoid cells, thin- to slightly thick-walled, up to 30 μm wide. Pileipellis a sometimes slightly loosening hymeniderm composed of 21.5–28 x 11.5–19 μm, clavate to vesiculose, thin- to slightly thick-walled, smooth, hyaline cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 10 μm wide hyphae. Caulocystidia absent; scattered adpressed to erect, clavate to cylindrical terminal cells present. Clamp-connections present in all tissues. – Fig. 51

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. On soil in a marshy forest and on dead leaves.

 

Distribution. Known only from the Democratic Republic of Congo and Uganda.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Equateur Province, Binga, Nov. 1925, M. Goossens–Fontana 397 (BR 11453–07, lectotype).

Uganda. Mpanga Forest, 30 March 1957, A. French 14 (K(M) 134417); Buganda Province, Mengo District, Mabira Forest, 9 June 1968, D.N. Pegler U 1373 (K(M) 134416).

 

Notes. Marasmius goossensiae is characterised by having rather large carpophores with a ± cream or grey cream pileus and concolorous lamellae and stipe; microscopically it has small basidiospores, clavate and rather voluminous cheilocystidia, well-developed large pleurocystidia and no caulocystidia.

Singer (1964, 1965a) mentioned narrower basidiospores (3.0–3.5 μm), and narrower basidia (4.0–7.0 μm). Pegler (1977) described a cream to pale orange pileus, fuscous brown at centre, a pale buff, soon darkening stipe, slightly smaller basidiospores (4.8–7.5 x 2.5–3.8 μm), smaller basidia (18–25 x 4–6 μm), larger cheilocystidia (36–50 x 9–13 μm), and smaller pleurocystidia (54–70 x 8.5–11 μm); these features may fall within the variability of this less known species.

Among tropical species with small basidiospores (less than 10 μm long) and well-developed pleurocystidia, M. witteanus Singer differs especially in having smaller and darker coloured carpophores, a thinner densely cespitose stipe (1–2 mm) and smaller basidiospores ((4.6–)5.0–6.2(–6.6) x (2.5–)3.1–3.8(–4.2) μm). Marasmius ochraceus Berk. & Broome seems to be very similar with a brightly tawny brown pileus pallescent to ochraceous towards margin and smaller basidiospores (5.5–7.2 x 2.7–3.5 μm); M. calvus Berk. & Broome has a campanulate pileus, larger and differently shaped basidiospores (8–10.5 x 2.7–3.5 μm, cylindrical to subfusoid) and smaller refractive cystidia (30–35 x 6–8 μm); both species are known from Sri Lanka (Pegler 1986). Marasmius phlebodiscus Desjardin & E. Horak, from Papua New Guinea, has a pale beige to tan coloured pileus with net-like wrinkles at centre, a densely longitudinally fibrillose stipe, smaller basidiospores (5–6.5 x 3–3.5 μm) and only fusoid pleurocystidia (Desjardin & Horak 1997). Marasmius riparius Singer, from Argentina, has a differently coloured pileus (like M. oreades), a white then cinnamomeous stipe, larger (8.2–10 x 4.8–5.5 μm) ellipsoid-fusoid basidiospores and smaller cystidia (40–44 x 8–9 μm); M. silvicola Singer has a larger (17–95 mm), deeply cinnamomeous to ochraceous brownish pileus, a larger stipe (20–118 x 1.5–7 mm), and cheilocystidia different from pleurocystidia (Singer 1976).

Marasmius goossensiae is known under the local names “lokombe” and “sensanse” (Hendrickx 1948).

 

48.  Marasmius muramwyanensis Antonín

Antonín, Mycotaxon 85: 124 (2003). Type: Burundi, Muramwya Province, Teza, 20 Dec. 1978, J. Rammeloo 6148 (BR 11905–71, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 124–126 (2003).

 

Pileus up to 45 mm broad, conical when young, later broadly conical, with small broad umbo, with regular, sometimes slightly undulate margin, glabrous, membranaceous, up to 1 mm thick, brown-beige (± 6E5) at centre, paler towards margin, beige or greyish orange (± 5B4). Lamellae close, ca. 35–40, l = 3, free, convex to convex-concave, up to 2 mm broad, not intervenose, pale yellow to yellowish white (3A2 to 4A2), with entire or weakly undulate concolorous edge. Stipe 40–65 x 3 mm, cylindrical, sometimes compressed, sometimes twisted, mealy to finely pubescent, finely fibrillose to very finely fibrillose-squamulose towards base, very pale, sordid white to cream at apex, ochraceous to brownish orange (5A4) at base. Context up to 1 mm thick in pileus, brownish under pileipellis, pale beige otherwise, with rather unpleasant terricolous smell and indistinct, after longer mastication fungoid taste— Pl. 8

 

Basidiospores 5.0–6.2 x 2.9–3.7 μm, E = 1.5–1.9, Q = 1.7, ellipsoid to broadly ellipsoid, both thin-walled non-dextrinoid and slightly thick-walled dextrinoid (crassospores), smooth. Basidia 27–28 x 5.8–6.2 μm, 4-spored, clavate; crassobasidia also present. Basidioles 11.5–27 x 2.5–7.0 μm, clavate, cylindrical. Cheilocystidia 10–23(–35) x 4.0–7.0 μm, clavate, (sub)cylindrical, subfusoid, sometimes septate, often irregular, thin-walled. Pleurocystidia 19–55 x 8.5–17 μm, variable, often voluminous, cylindrical, clavate, fusoid, often subrostrate, hyaline, thin-walled, with refractive contents. Hyphae cylindrical to subinflated, thin- to slightly thick-walled (often in subpileipellis), pale yellowish, up to 12 μm wide. Pileipellis a hymeniderm composed of 16–35 x (7.0–)10.0–19 μm, clavate, smooth, thin- to slightly thick-walled cells, with yellowish to brownish walls in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, incrusted, up to 7.0 μm wide hyphae. Caulocystidia 22–42 x (4.0–)8.0–13 μm, adpressed to erect, clavate, cylindrical, subfusoid, obtuse, thin- to slightly thick-walled. Clamp-connections present in all tissues. – Fig. 52

 

Chemical reactions. All hyphae and crassospores dextrinoid, other structures non-dextrinoid.

 

Ecology. Gregarious, on dead wood of cultivated Cupressus.

 

Distribution. Known only from the type locality in Burundi.

 

Revised specimens from tropical Africa.

Burundi. Muramwya Province, Teza, 20 Dec. 1978, J. Rammeloo 6148 (BR 11905–71, holotype).

 

Notes. Marasmius muramwyanensis is characterised by having a moderately large pileus, close lamellae, a finely pubescent stipe, small basidiospores, pleurocystidia which differs from cheilocystidia, and well-developed caulocystidia.

Marasmius witteanus Singer and M. goossensiae Beeli seem to be macroscopically close to this species. However, M. witteanus differs especially by a smaller pileus, a densely cespitose stipe, pleurocystidia similar to cheilocystidia and by the absence of caulocystidia; M. goossensiae has a differently coloured pileus, larger basidiospores, larger cheilo- and pleurocystidia, and no caulocystidia. Marasmius ochraceus Berk. & Broome has a large pileus (30–80 mm) and a thicker stipe (60–110 x 4–7 mm); M. calvus Berk. & Broome has a campanulate pileus, larger differently shaped basidiospores (8–10.5 x 2.7–3.5 μm, cylindrical to subfusoid) and smaller refractive cystidia (30–35 x 6–8 μm); both species are known from Sri Lanka (Pegler 1986). Marasmius phlebodiscus Desjardin & E. Horak, from Papua New Guinea, has a pale beige to tan coloured pileus with net-like wrinkles at centre, a densely longitudinally fibrillose stipe, and only fusoid pleurocystidia (Desjardin & Horak 1997). Marasmius riparius Singer, from Argentina, has a differently coloured pileus (like M. oreades), a white then cinnamomeous stipe, larger (8.2–10 x 4.8–5.5 μm) ellipsoid-fusoid basidiospores and smaller cystidia (40–44 x 8–9 μm) (Singer 1976). Marasmius silvicola Singer has a larger (17–95 mm), deeply cinnamomeous to ochraceous brownish pileus, a larger stipe (20–118 x 1.5–7 mm), and larger basidiospores (4.5–8.3 x 2.5–4.3 μm) (Singer 1976). 

 

49. Marasmius flavus Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 344 (1964). Type: Democratic Republic of Congo, Equateur Province, Eala, May 1923, M. Goossens–Fontana 151 (BR 11450–04, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 344 (1964); Singer, Flore Icon. Champ. Congo 14: 262 & Pl. 44, fig. 2 (1965).

 

Pileus 31–34 mm broad, convex, strongly sulcate, with undulate margin, glabrous, yellow to dirty yellow. Lamellae distant, L = ca. 15–17, l = 1–2, almost free or adnexed, often emarginate, rather broad, yellowish. Stipe 80 x 2–2.5 mm, subcylindrical or slightly broadened at base, hollow, brown-red; with developed basal mycelium. Spore print white. (According to Singer (1964, 1965a)). — Pl. 8

 

Basidiospores (only two basidiospores found) 13.5–15 x 3.1–3.5 μm, clavate, cylindrical-clavate, smooth, hyaline. Basidia not observed. Basidioles 30–39 x 4.0–11 μm, cylindrical to clavate. Cheilocystidia 28.5–46 x 13–18.5 μm, clavate, ± thin-walled, hyaline. Pleurocystidia 40–77 x 11.5–20(–23) μm, clavate to subfusoid, often subrostrate, thin-walled, hyaline. Hyphae cylindrical to subinflated, thin- or slightly thick-walled, up to 12(–17) μm wide. Pileipellis a hymeniderm composed of 17–27 x 11.5–16 μm, clavate to (sub)vesiculose, smooth, thin- to slightly thick-walled cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 7.0 μm wide hyphae. Caulocystidia absent; scattered terminal cells present on stipe surface. Clamp-connections present in all tissues. – Fig. 53

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, on litter in a marshy forest.

 

Distribution. So far known only from the Democratic Republic of Congo.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Equateur Province, Eala, May 1923, M. Goossens–Fontana 151 (BR 11450–04, holotype); ? Tshopo Province, close to Kisangani, 14 Apr. 1984, B. Buyck 1398 (BR 11759–22).

 

Notes. Marasmius flavus is characterised macroscopically in having a moderately large yellow pileus and a rather long red-brown stipe, microscopically especially by developed pleurocystidia, clavate, rather broad cheilocystidia and rather large basidioles. I found only a few basidiospores in the revision of the type specimen.

Of other tropical Marasmius species with yellow coloured carpophores, M. caryotae (Berk.) Petch, from Sri Lanka, has larger basidiospores (19–25 x 5–6.2 μm), larger basidia (40–45 x 9–10 μm), small cheilocystidia (15–20 x 10–12 μm) and no pleurocystidia (Pegler 1986; Petch 1948); M. rhyssophyllus Mont., from the Lesser Antilles, has a non-sulcate pileus, a small stipe (10–30 x 1–1.5 mm), small basidiospores (5–6.5 x 3.5–4.5 μm, however, 7–8.3 x 3.3–3.8 μm according to Singer (1976), and 6–7 x 4–4.5 according to Dennis (1970)), smaller basidia (22–28 x 5–6 μm) and no pleurocystidia (Pegler 1983).

A description by Singer (1964, 1965a) differs by the size of the pleurocystidia (30–43 x 8.0–14.5 μm) and smaller pileipellis cells (about 16 x 11.5–12.5 μm); he also found only a few basidiospores.

Beeli (1928a) included the type specimen (Goossens–Fontana 151) in M. bekolacongoli var. brunneolus Beeli [= M. brunneolus (Beeli) Singer]. However, M. brunneolus differs especially in having larger carpophores, a fuligineous brown, often striped pileus, larger basidiospores (15.5–25.5 x 3.8–5.4 μm), and by the absence of pleurocystidia.

Collection B. Buyck 1398 represents a microscopically very similar fungus. Macroscopically it has a darker pileus (greyish orange to yellowish brown) when young, then light yellow at centre and greyish, greyish yellow, light yellow to yellow sulci and brownish yellow to yellowish brown striae. However, pileus is yellowish brown (not yellow to dirty yellow) in the type drawing of M. flavus by Goossens–Fontana (Singer 1965a). Therefore, the collection by Buyck may belong here.

  

50. Marasmius heinemannianus Antonín

Antonín, Belg. Journ. Bot. 131(2): 127 (1998).  Type: Benin, Atacora Province, Boukombé, 25 Aug. 1997, Antonín B97.101 (BR 101146–72, holotype; BRNM 612544, isotype).

 

Selected descriptions and icons. Antonín, Belg. Journ. Bot. 131(2): 127–131 (1998).

 

Pileus 6–80 mm broad, hemispherical-conical, without central umbo, slightly involute at margin when young, later conical-convex, mostly with slightly pronounced obtuse broad to truncate umbo, and slightly inflexed to more or less straight margin, then almost applanate with low broad central papilla, rarely applanate without papilla or with a small central depression when old, hygrophanous, rarely (old specimens) slightly translucently striate, very finely tomentose-pruinose when young, then more or less glabrous (except for finely tomentose-pubescent margin), smooth, later on finely rugulose, sometimes cracking, sometimes with concentrical rugulose zones (like Russula olivacea), not striate or finely rugulose-striate (when old) at margin, undulate when old, pileipellis sometimes longer than lamellae; reddish-brown (8B–E8, 9D7) at centre, and brownish red (more yellow than 8C7) towards margin when young, becoming darker reddish brown (up to 9E7) at centre and distinctly paler yellow-orange (5–6A6–7), sometimes even up to reddish yellow (4–5A4–5) towards margin, some parts almost whitish-yellowish. Lamellae distant, L = 22–35, l = 2–3, emarginate and attached by small tooth, rather thick, more or less horizontal when young, ventricose (up to 20 mm broad) when old, not intervenose or intervenose only when old; almost white when young, pale yellow (3–4A2–3) when old; edge concolorous, entire, then uneven to serrulate, finely pubescent. Stipe 27–60 x 4–12 mm, cylindrical, slightly broadened at apex, slightly tapered, cylindrical or slightly clavate towards base, sometimes laterally compressed, rarely with a groove, in older specimens longitudinally striate-sulcate, often slightly curved, sometimes twisted, entirely finely tomentose, with whitish woolly tomentum at base; white to pale cream or orange-cream (paler than 3A2) when young, then darker, orange to brownish orange (5A3–4, 6A6 to 7B–C6), almost white to whitish towards apex; forming dirty whitish to reddish rhizoids at base. Context white, concolorous with surface under pilei- and stipitipellis, cartilaginous and thin (up to 3 mm) in pileus, hollow, tough-elastic in stipe (not so much as M. oreades); smell (acid) fungoid, slightly unpleasant, taste mild, after longer mastication sometimes slightly astringent. Spore print pale yellowish white (2A2–4A2). — Pl. 9

 

Basidiospores 10.8–14.5 x (4.2–)4.6–6.0(–6.6) μm, E = 1.8–2.5, Q = 2.1–2.5(–2.9), ellipsoid or subfusoid, hyaline, smooth, mostly thin-walled and non-dextrinoid; slightly thick-walled and dextrinoid crassospores always present, rare or numerous, with or without a large central vacuole. Basidia 25–38.5 x 8.1–11.0 μm, 4-spored, clavate; dextrinoid crassobasidia also present. Basidioles 14–31 x 3.0–9.0 μm, clavate to cylindrical, less frequently subfusoid, thin-walled. Cheilocystidia (16–)22–61.5 x 6.2–12.5 μm, very variable in form, cylindrical, clavate, fusoid, irregular, sometimes rostrate, rarely branched, thin-walled, hyaline, more distinct when old. Pleurocystidia 27–46.5 x 5.5–11.5 μm, narrowly clavate, subfusoid, rarely sublageniform or subcylindrical, thin-walled, non-dextrinoid, sometimes slightly refractive in KOH. Hyphae cylindrical to inflated, thin- to slightly thick-walled, smooth or incrusted, branched, hyaline, incrusted hyphae pale orange-brownish, 5.0–23 μm wide. Pileipellis a hymeniderm, sometimes slightly disintegrated when old, composed of 17.5–54 x (7.0–)9.0–17 μm, clavate, narrowly clavate, subcylindrical, subfusoid, sometimes subrostrate or subcapitate, sometimes slightly irregular, thin-, less frequently slightly thick-walled, pale orange-brownish to subhyaline cells. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, finely incrusted, 2.0–6.2 μm wide hyphae, with pale orange coloured walls in KOH. Caulocystidia 15.5–42 x (4.6–)5.1–11 μm, cylindrical to narrowly clavate or subfusoid, obtuse, thin- or slightly thick-walled, hyaline to pale orange, often forming groups. Clamp-connections abundant in all tissues.  Fig. 54

 

Chemical reactions. All hyphae, crassobasidia and crassospores dextrinoid, other structures non-dextrinoid.

 

Ecology. Single or in groups on soil, on open places in humid grassland and on pastures. 

 

Distribution. Known only from some localities in Benin.

 

Revised specimens from tropical Africa.

Benin. Atacora Province, Boukombé, 25 Aug. 1997, V. Antonín B97.101 (BR 101146–72, holotype; BRNM 612544, isotype); Ibid., SE of town, 1 Sept. 1997, V. Antonín B97.140 (BR 101184–13); Atacora Province, Kounagnigou, 3 Sept. 1997, V. Antonín B97.149 (BR 101191–20); Atacora Province, Kota, 27 Aug. 1997, V. Antonín B97.118 (BR 101163–89).

 

Notes. Marasmius heinemannianus is characterised by having rather large, fleshy carpophores, reminiscent of the habitus of M. oreades, a brightly orange-brown, reddish-brown to yellow-orange pileus, a yellowish white spore print, rather large basidiospores, well-developed cheilo-, pleuro-, as well as caulocystidia, and its habitat - it grows on open grassy places.

From the African species of this section bearing pleurocystidia, Marasmius flavus Singer differs especially in having a smaller, yellow coloured, sulcate-striate pileus, a long and narrow stipe (up to 80 x 2–2.5 mm), narrower basidiospores (13.5–14.6 x 3.1–3.5 μm), and broader pleurocystidia (42.5–69 x 11.5–19 μm); Marasmius goossensiae Beeli differs especially by its brown to greyish, striate pileus, smaller basidiospores (6.2–7.9 x 3.5–4.2 μm), and broader pleurocystidia (47.5–92.5 x 12–23 μm); Marasmius witteanus Singer has a smaller (8–18 mm broad), ochraceous brown pileus, a long and narrow cespitose stipe (50–110 x 1–2 mm), smaller basidiospores (5.4–6.6 x 3.1–4.0 μm), and larger pleurocystidia [37–78 x 10–23(–34) μm]. Marasmius titanosporus Reid & Guillarmod, described from South Africa (Reid & Guillarmod 1988), which may also have developed pleurocystidia, differs especially in having a distinctly sulcate, pale grey to blackish grey pileus, distant lamellae (L = 15–18), a long and narrow stipe (up to 90 x 6 mm), and larger basidiospores (22–36 x 5.0–7.2 μm).

From the species with pleurocystidia from North America (Gilliam 1976, Halling 1983), M. cystidiosus (A.H. Sm. & Hesler) Gilliam differs especially in having a yellowish pink, yellowish brown to orange-yellow coloured and when moist translucent pileus, a stipe without caulocystidia, a bitter taste, and smaller basidiospores (7.6–10.5(–12.5) x 2.8–3.9 μm). Marasmius leighii A.H. Sm., which is sometimes considered identical with the previous species, lacks caulocystidia, and grows cespitosely on decaying hardwood. Marasmius lilacinus (Coker & Beardslee) Singer has a tan coloured pileus with lilac tinge when old, a long and narrow stipe (up to 110 x 6.5 mm), white spore print, smaller basidiospores (6.5–8.0 x 4.0–5.0 μm), and larger pleurocystidia (45.0–60.0 x 13.0–23.0 μm). Marasmius nigrodiscus (Peck) Halling, has a coloured, buff or olivaceous spore print, an umbrinous, towards the margin paler pileus, smaller basidiospores ((5.0–)7.0–9.0 x 4.0–5.0 μm), larger pleurocystidia (45–120 x 7.0–18 μm), and grows mostly in forests.

Marasmius plumieri (Lév.) Singer s. Pegler, originally described from the Caribbean area, differs especially in having a deeply plicate-striate pileus, brownish lamellae, narrower basidiospores (14.5–15.5 x 4.0–4.5 μm) and no cheilo- and pleurocystidia (Pegler 1983). Another South-American species, M. fiardii Singer, differs especially by its dark brown to greyish, plicate-striate pileus, larger basidiospores (13.5–19.5 x 3.5–4.6 μm), the lack of pleurocystidia, and differently shaped pileipellis cells (Pegler 1983). Among the species published by Singer (1976) from South America, Marasmius viegasii Singer differs especially by its sulcate pileus, a white spore print, narrower basidiospores (16 x 4.0 μm), the absence of cheilo- and pleurocystidia, and a different ecology (it causes root rot of the coffee tree in Brazil); M. myocephalus Singer is smaller, with a mouse grey pileus, smaller basidiospores ((6.0–)10.5–11.5 x 3.0–4.0 μm), lacking cheilo- and pleurocystidia, and with a different ecology; it grows on leaves and roots in tropical montane forests. Among the South-American species bearing pleurocystidia, Marasmius riparius Singer, coloured like M. oreades, differs by a deeply sulcate pileus, distinctly smaller basidiospores (8.2–10.0 x 4.8–5.5 μm) and lacking cheilocystidia. Marasmius silvicola Singer has a striate, cinnamon to ochraceous-brownish pileus, a greyish stipe and smaller basidiospores (4.5–8.3 x 2.5–4.2 μm).

Stevenson (1964) published from New Zealand only M. oreades from this group. Also no macroscopically and microscopically similar taxon has been published in the Marasmius monograph of Papua New Guinea, New Caledonia, and New Zealand species (Desjardin & Horak 1997). Marasmius maximus Hongo from Japan, mentioned as close to M. oreades (Hongo 1962), differs especially in having a sulcate-striate, ochraceous-alutaceous coloured pileus, and smaller basidiospores (7.0–9.0 x 3.0–4.0 μm). Another Japanese species, M. aurantioferrugineus Hongo, having an up to 70 mm broad, orange-ferrugineous pileus seems to be very close. According to the original description (Hongo 1965), it differs especially by its longer and narrower white stipe (50–120 x 3–6 mm) with bulbous base and narrower basidiospores (11–12 x 4.5 μm). Hongo (1962, 1965) did not mention the presence of pleurocystidia. M. brunneospermus Har. Takah., described also from Japan, differs especially in having a brown spore print (6E4–6E6) mottled with white parts, a less brightly coloured pileus, smaller basidiospores (6.4–8.0 x 2.4–3.6 μm), and larger pleurocystidia (50–95 x 5.0–13.5 μm). Especially its spore print colour represents a unique feature in the genus Marasmius. Pegler (1986) published two species with fleshy carpophores and present pleurocystidia from Sri Lanka. Marasmius ochraceus Berk. & Broome has a tawny brown to ochraceous pileus, lamellae becoming umbrinous when dry, distinctly smaller basidiospores (5.5–7.2 x 2.7–3.5 μm), and grows on rotten wood. Marasmius calvus Berk. & Broome is smaller (pileus 20–45 mm broad), pale fawn to fuscous brown, with small basidiospores (8.0–10.5 x 2.7–3.5 μm), and grows on wooden sticks. Corner (1996) described M. trogioides Corner with an up to 90 mm broad cinnamon fawn to fulvous ochraceous or fuscous brown pileus, smaller basidiospores (9.0–11.5 x 4.7–5.5 μm), larger cheilo- and pleurocystidia different in shape and clampless hyphae, and M. benecystidiatus Corner with a smaller, 18–40 mm broad, rugoso-reticulate, dark brown to fawn ochraceous pileus, smaller basidiospores (9.0–12.5 x 3.0–3.5 μm), and larger fusoid-subventricose thick-walled cystidia.

 

51. Marasmius flavidulus Henn.

Hennings, Bot. Jahrb. Syst. 30: 49 (1901). Type: Cameroon, Bipinde, virgin forest area (Urwaldgebiet), 1900, G. Zenker, in: G. Zenker, Flora von Kamerun No. 2198 (K(M) 92586, syntype).

 

Selected descriptions and icons. Hennings, Bot. Jahrb. Syst. 30: 49 (1901).

 

Pileus 20–30 mm broad, membranaceous, campanulate-applanate, with umbilicate or depressed centre, papillate, radially striate or subsulcate, yellowish (flavidulus). Lamellae adnate, subdistant, ventricose, 2 mm broad, concolorous with pileus and stipe. Stipe 60–100 x 1–1.5 mm, fistulose, thin-coriaceous, pruinate, yellowish (flavidulus). (According to Hennings 1901).

 

Basidiospores 26–30 x 5.0–6.0 μm (only six found), narrowly clavate, lacrimoid, sometimes curved, thin-walled, hyaline. Basidia 37–40 x 8.5–9.0 μm, clavate. Basidioles 22–40 x 5.0–10.0 μm, clavate, cylindrical, (sub)fusoid. Cheilocystidia 16–23 x 9.0–12 μm, clavate to subfusoid, thin-walled. Pleurocystidia numerous, 30–50 x (5.0–)8.0–19 μm, fusoid, clavate, thin-walled. Trama hyphae cylindrical to (sub)inflated, thin-walled, (sub)hyaline in KOH, up to 20 μm wide. Pileipellis a hymeniderm composed of 22–28(–45) x 10–20 μm, (broadly) clavate, pyriform, sometimes subvesiculose, thin- to slightly thick-walled cells, with subhyaline to pale greyish-yellowish walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 6.0 μm wide hyphae with pale yellowish to ochraceous walls in KOH. Caulocystidia 22–57 x 5.0–10.0 μm, adpressed to erect, cylindrical, clavate, fusoid, sometimes rostrate, thin-walled, dextrinoid. Clamp-connections present in all tissues. – Fig. 55

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. In a virgin forest area (Urwaldgebiet).

 

Distribution. Known only from Cameroon.

 

Revised specimens from tropical Africa.

Cameroon. Bipinde, 1900, G. Zenker, in: G. Zenker, Flora von Kamerun, No. 2198 (K(M) 92586, syntype).

 

Notes. Marasmius flavidulus is characterised by having a yellowish pileus, large basidiospores, small clavate cheilocystidia and by the absence of caulocystidia.

Close species seems to be M. flavus Singer, however, with distinctly smaller basidiospores and larger cheilo- and pleurocystidia. Marasmius caryotae (Berk.) Petch with a straw yellow to greyish yellow pileus has smaller basidiospores (16–25 x 5–6.2 μm) and no pleurocystidia are developed (Pegler 1986). Among species with large basidiospores, M. amabilis Hariot & Pat. has a white to cream buff pileus; M. musisporus Desjardin & E. Horak even larger basidiospores (30–40 x 4.5–5 μm) and a greyish lilac pileus disc and furrows, and M. purpureostriatus Hongo has a purple pileus disc and furrows; moreover no pleurocystidia are developed in any of these species (Desjardin & Horak 1997).

  

52. Marasmius latepileatus Antonín & C. Sharp

Antonín & C. Sharp, Mycotaxon 88: 58 (2003). Type: Zimbabwe, Kwekwe, Sagle Park, 1829D4, 27 Febr. 1981, Sharp 817/97 (BR, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 58–60 (2003).

 

Pileus up to 100 mm broad, campanulate, then convex to applanate, glabrous, with greasy appearance, except for smooth centre deeply sulcate, beige cream with darker centre. Lamellae distant, L = 20, l = 2, sinuate, adnate or free, ventricose, up to 12 mm broad, cream coloured. Stipe up to 140 x 8 mm, cylindrical, slightly laterally compressed, fistulose, thin-coriaceous, smooth, glabrous, beige cream at apex, up to tan to golden brown towards base, with white basal mycelium. Smell unpleasant, of bad potatoes.

 

Basidiospores 23–30 x 5.0–7.5 μm, E = 3.7–4.5, Q = 4.3, narrowly clavate, lacrimoid, sometimes curved, thin-walled, hyaline. Basidia 34–38 x 8.0–11.0 μm, 4-spored, clavate. Basidioles 25–40 x 5.0–10.0 μm, clavate, cylindrical, (sub)fusoid. Cheilocystidia 12–25 x 7.0–11 μm, (broadly) clavate to subfusoid, thin-walled. Pleurocystidia numerous, 37–60 x 9.0–15 μm, fusoid, clavate, thin-walled, with refractive contents. Trama hyphae cylindrical to (sub)inflated, thin-walled, (sub)hyaline in KOH, up to 12 μm wide. Pileipellis a hymeniderm of 18–30 x 9.0–16 μm, (broadly) clavate, pyriform, ± thin-walled cells, with subhyaline to pale yellowish walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, smooth, up to 5.0 μm wide hyphae with pale ochraceous walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 56

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing in dense clusters or in large groups in a miombo woodland. In humus and leaf litter of Tarenna neurophylla, Brachystegia boehmii and Securinega virosa, under Strychnos potatorum, Tarenna neurophylla, Dichrostachys cinerea, Securinega virosa, Brachystegia boehmii, Aloe excelsa and Grewia praecox.

 

Distribution. Known only from the type locality in Zimbabwe.

 

Revised specimens from tropical Africa.

Zimbabwe. Kwekwe, Sagle Park, 1829D4, 27 Febr. 1981, Sharp 817/97 (BR, holotype).

 

Notes. Marasmius latepileatus is characterised by having very large carpophores, a beige cream pileus with darker centre, very broad lamellae, large basidiospores, small clavate cheilocystidia, well-developed pleurocystidia and by the absence of caulocystidia. It is not edible and its local name is “chinyakacheche” (Shona) (Sharp in litt.).

Marasmius flavidulus Henn. has a similar size of basidiospores and colour of carpophores, However, it has distinctly smaller carpophores and well-developed caulocystidia. Marasmius bekolacongoli Beeli and M. brunneolus (Beeli) Singer also have very large carpophores. Both have a differently coloured pileus and lack pleurocystidia. Among species growing outside of tropical Africa, no species with similar combination of macroscopic and microscopic features has been found in Corner (1996), Dennis (1970), Desjardin & Horak (1997) or Pegler (1977, 1983, 1986).

 

53. Marasmius albidocremeus Antonín

Antonín, Mycotaxon 85: 110 (2003). Type: Cameroon, Dja Biosphere Reserve, near Somalomo, 7 April 2001, V. Antonín Cm 01.27 (BRNM 666094, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 110–111 (2003).

 

Pileus 12–30 mm broad, broadly conical with truncate to obtuse centre, then plano-conical with obtuse to applanate centre, straight to uplifted at crenulate margin, except for rugulose centre, entirely distinctly striate-plicate, hygrophanous, translucent, whitish to pale cream (4A2–5A3) at centre, almost white when dried out, yellowish grey (slightly paler than 4B2) towards margin. Lamellae distant, L = 11–13, l = 2–3, slightly emarginate and ± adnate, ventricose, sometimes forming an adnexed false collarium, not intervenose, rather broad (up to 2.5 mm), cream, with concolorous edge. Stipe 50–90 x 0.5–1.25 mm, cylindrical, slightly broadened above, cylindrical to subclavate at base, glabrous, smooth, fistulose, whitish or cream at apex, orange brown to (orange) golden (6C7, 6B–D8) at base, with pale ochraceous basal mycelium. — Pl. 9

 

Basidiospores 16.5–23 x 3.5–5.0 μm, E = 3.8–5.7, Q = 4.6, narrowly clavate, sublacrimoid, sometimes curved, thin-walled, hyaline. Basidia 4-spored. Basidioles 12–29 x 3.0–8.0 μm, cylindrical, clavate, subfusoid. Cheilocystidia 10–31 x 6.0–11 μm, clavate, sometimes slightly irregular to lobate, smooth or with scattered projection(s), thin-walled. Pleurocystidia (21–)27–48 x (4.5–)6.5–12 μm, clavate, fusoid, subcylindrical, obtuse to subrostrate, hyaline. Hyphae cylindrical, thin-walled, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of 15–26 x 8.0–13(–15) μm, clavate to pyriform, thin- to slightly thick-walled, smooth cells, rarely irregular or with single projection(s), with subhyaline to pale yellowish walls in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 5.0 μm wide hyphae, with pale yellowish walls in KOH. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 57

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single or in groups, on strongly decayed wood, in a Gilbertiodendron dewevrei stand.

 

Distribution. Known from two localities in Cameroon.

 

Revised specimens from tropical Africa.

Cameroon. Dja Biosphere Reserve, near Somalomo, 7 April 2001, V. Antonín Cm 01.27 (BRNM 666094, holotype); Ibid., 8 April, V. Antonín Cm 01.51 (BRNM 666117).

 

Notes. Marasmius albidocremeus is characterised by having a whitish or pale cream to dirty yellowish grey coloured pileus, a rather long and slender stipe, large narrowly clavate, sublacrimoid, sometimes slightly curved spores, and by the presence of clavate cheilocystidia, clavate or fusoid pleurocystidia, and the absence of caulocystidia. It belongs to sect. Globulares Kühner.

Concerning species with well-developed pleurocystidia, Marasmius flavus Singer (holotype BR!) seems to be a close species; it differs only microscopically by smaller spores (13.5–15 x 3.1–3.5 μm), longer basidioles 30–39 x 4.0–11 μm, cheilocystidia (28.5–46 x 13–18.5 μm) and pleurocystidia (40–77 x 11.5–20(–23) μm).

 

54. Marasmius staudtii Henn.

Hennings, Bot. Jahrb. Syst. 22: 97 (1895).

 

54.1. M. staudtii var. staudtii

Type: Cameroon, Yaoundé, 15 March 1894, G. Zenker & Staudt 275 (not preserved).

 

Selected descriptions and icons. Hennings, Bot. Jahrb. Syst. 22: 97 (1895); Morris, Kirkia 13(2): 341 (1990) (as M. lilacinoalbus); Singer, Bull. Jard. Bot. Etat Brux. 34: 347–348 (1964); Singer, Flore Icon. Champ. Congo 14: 264 (1965).

 

Pileus 15–70 mm broad, campanulate, then campanulate-convex to plano-conical, with subumbonate to almost applanate centre, crenulate at margin, strongly plicate-sulcate, slightly rugulose when old, glabrous, whitish, cream to yellowish (3A2) at centre and sulci, with deeply purple (14F7–8) stripes. Lamellae distant, L = 8–16, l = 2–3, emarginate and shortly to broadly adnate, not intervenose or intervenose when old, broad, greyish whitish to greyish yellowish (e.g. 6B2), with concolorous edge. Stipe 50–150 x 1–3.5 mm, cylindrical or slightly broadened at base, hollow, smooth to slightly fibrillose, glabrous, sometimes twisted, white with a very pale violaceous tinge at apex, through a short brown (8D6) zone, dark brown (8E–F7) to black-brown (9F7) towards base; basal mycelium rich, tomentose, dirty white, a transient to membranaceous layer penetrates the surrounding substrae. — Pl. 9

 

Basidiospores (20–)24.5–28.5 x (4.5–)5.5–6.0 μm, E = (4.0–)4.5–5.4, Q = 4.7, clavate, hyaline. Basidia 33–44 x 8.5–10.5 μm, 4-spored, clavate. Basidioles 19–36(–42) x 2.5–11 μm, cylindrical, clavate, subfusoid. Cheilocystidia 15–26 x 7.5–11 μm, clavate, thin- to slightly thick-walled. Pleurocystidia 17–50 x 8.5–19 μm, clavate, fusoid, vesiculose, sometimes subrostrate, thin-walled, with slightly refractive contents, hyaline, sometimes less distinct. Hyphae cylindrical, thin-, sometimes slightly thick-walled, hyaline, slightly brownish in subpileipellis, 2.5–10.5 μm wide. Pileipellis a hymeniderm composed of 16–31 x 10–14 μm, clavate to vesiculose, sometimes capitate, thin- to slightly thick-walled, smooth cells, with pale brownish walls in KOH. Stipitipellis a cutis consisting of parallel, cylindrical, slightly thick-walled, up to 5 μm wide hyphae, olivaceous greenish to olivaceous brownish in KOH. Caulocystidia absent. Clamp-connections abundant in all tissues. – Fig. 58

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, on decaying fallen leaves, and on twigs in humus layer, in a primary rain forest with Gilbertiodendron dewevrei.

 

Distribution. Known only from Cameroon (Hennings 1895, Singer 1964, 1965a), the Democratic Republic of Congo, Ethiopia, Kenya, Malawi and Zambia. Outside of tropical Africa, this species was recorded from Guandong and Hainan Province in China (Li Taihui & al. 1990).

 

Revised specimens from tropical Africa.

Cameroon. Dja Biosphere Reserve, ca. 2–3 km South of Somalomo, 7 April 2001, V. Antonín Cm 01.35 (BRNM 666104); Ibid., 7 April 2001, D.C. Mossebo 295 (herb. Mossebo).

Democratic Republic of Congo. Kisantu, 20 March 1907, H. Vanderyst s. n. (BR 11507–61); Kimpako, 18 Dec. 1908, H. Vanderyst s. n. (BR 11508–62); Kivu Province, Irangi, J. Rammeloo Z 461 (GENT); Ibid., 26 March 1972, J. Rammeloo Z 183 (GENT); ? Tshopo Province, close to Batiabongena, Km 14 on the road to Buta, 16 April 1984, B. Buyck 1443 (BR 11754–17); ? Tshopo Province, close to Kisangani, Isle of Congolo, 14 April 1984, B. Buyck 1396 (BR 11761–24).

Ethiopia. Shoa Province, Wondo Genet Forestry College near Shashemene, 18 July 1990, L. Ryvarden 28042 (K(M) 16362, as M. bekolacongoli).

Kenya. Nyanza Province, Kericho District, Kigumu River, Kericho, 25 March 1968, D.N. Pegler K 229 (K(M) 8828, as M. bekolacongoli); Eastern Province, Aberdare Mts., Kimakia Forest station, 16–18 Jan. 1973, L. Ryvarden 9059 (K(M) 8830, as M. lilacinoalbus).

Malawi. Mulanje, Ruo Estate, 19 Dec. 1981, B. Morris 400 (K(M) 8829, as M. lilacinoalbus).

Zambia. Kitwe, 7 Dec. 1978, G.D. Piearce FP 593/7 (K(M) 134373); Ndola, 18 Dec. 1980, G.D. Piearce FP 678/1 (K(M) 111250, as M. bekolacongoli).

 

Notes. Marasmius staudtii is characterised by having a rather small, distinctly sulcate, violet and cream striped pileus, a rather long stipe, large basidiospores, well-developed pleurocystidia, stipitipellis cells olivaceous tinged in KOH, and lacking caulocystidia. However, only a few basidiospores and basidia were found in the revised specimens.

A combination of the presence of pleurocystidia and violaceous colour is not common in marasmioid fungi. According to Desjardin & Horak (1997), Marasmius staudtii may represent an older name for M. purpureostriatus Hongo. In comparison with the description (Desjardin & Horak 1997, isotype ZT!), the last species differs only in having smaller cheilocystidia (15–37 x 10.0–15 μm), and totally different hosts (trees of the family Fagaceae).

Singer (1964, 1965a) did not mention the presence of pleurocystidia, and described slightly narrower basidiospores (x 3.7–5.0 μm). Hennings (1985) described Marasmius staudtii var. pallida Henn. (Cameroon, 15 March 1894, Zenker & Staudt 276) with a pale greyish pileus and a yellow-brown stipe. Singer (1964, 1965a) synonymised it (with a question mark) with M. brunneolus (Beeli) Singer.

Collections by J. Rammeloo (JR Z183) and B. Buyck (BB 1443) differ from the typical specimens of var. staudtii in having smaller (16–22 x (4.0–)4.5–5.5 μm, E = 3.2–4.6, Q = 3.7) basidiospores (the typical var. staudtii: (20–)24.5–28.5 x (4.5–)5.5–6.0 μm, E = (4.0–)4.5–5.4, Q = 4.7). Other macroscopic and microscopic features agree very well. For the time being, I consider this spore size as belonging to the variability of this species; this decision is supported by the fact, that both types were collected in the same locality. Moreover, in herbarium specimen J. Rammeloo Z183, two types of basidiospores were found: basidiospores from lamellae measuring 16–22 x (4.0–)4.5–5.5 μm, basidiospores from an included spore print 24–30 x 7.0–9.0 μm.

 

54.2. M. staudtii var. magnisporus Antonín Antonín, Mycotaxon 88: 68 (2003). Type: Democratic Republic of Congo, Kivu Province, Irangi, April 1972, J. Rammeloo Z 390 (GENT, holotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 88: 68–69 (2003).

 

Pileus 22–42 mm broad, convex to campanulate, strongly sulcate, crenulate at margin, glabrous, membranaceous, always in violaceous colour, reddish grey, brownish grey to dull red (11B2, 11C2–3), greyish magenta to dark purple (14D–F5–6) at centre and sulci, yellowish white to pale yellow (3A2–3) or pale violaceous (greyish ruby, 12B–C3) in striae. Lamellae distant, L = ca. 18–30, l = 1–2, narrowly adnate to free, not intervenose, yellowish white (3A2), with concolorous, entire edge. Stipe 60–130 x 1–2 mm, cylindrical or compressed, slightly broadened at base (2 mm), glabrous, hollow, twisted-grooved in upper part, black-brown towards base, paler, brown (7D6) at apex; with a rich basal mycelium. Context white, with indistinct smell and slightly bitter taste.

 

Basidiospores 32–44 x 5.5–7.5 μm, E = 4.8–6.8, Q = 5.9, narrowly clavate, narrowly fusoid, thin-walled, hyaline. Basidia 27–41 x (8.5–)10–17 μm, 4-spored, clavate or subutriform. Basidioles 18.5–45.5 x 4.4–17 μm, clavate, cylindrical. Cheilocystidia 24–35(–42.5) x (8.5–)10–13.5(–17) μm, clavate, often capitate, thin-, rarely slightly thick-walled. Pleurocystidia 35–50 x (8.5–)12–25(–28) μm, broadly clavate, clavate, vesiculose to fusoid, thin-walled, with refractive contents. Hyphae of cylindrical, subinflated to fusoid cells, branched or with lateral projections (especially in pileus), thin- to slightly thick-walled, up to 15.5 μm. Pileipellis a hymeniderm composed of 17–35 x (9.2–)12–19 μm, clavate to vesiculose, sometimes pyriform, thin- to slightly thick-walled, smooth cells, with subhyaline to pale greyish brownish walls in KOH; sometimes with more pileipellis cells on one ± (sub)globose, sometimes irregular cell in subpileipellis. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 7.0 μm wide hyphae, with olivaceous green walls in KOH. Caulocystidia absent; scattered adpressed to erect, cylindrical to clavate terminal cells present. Clamp-connections present in all tissues. – Fig. 58

 

Chemical reactions. All hyphae and pileipellis cells dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, probably on twigs in a dense leaf humus layer, in a primary rain forest.

 

Distribution. Known only from the Democratic Republic of Congo and Zambia.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Kivu Province, Irangi, April 1972, J. Rammeloo Z 390 (holotype, GENT); Ibid., J. Rammeloo  Z 390a (GENT).

Zambia. Kitwe, 7 Dec. 1978, G.D. Piearce FP 593/6 (K(M) 111242, as M. zenkeri); Ibid., 14. Dec. 1978, G.D. Piearce FP 596/2 (K(M) 111243, as M. zenkeri).

 

Notes. This variety is characterised by having a violaceous to dark purple pileus with pale stripes, very large basidiospores, rather small pleurocystidia and stipitipellis hyphae olivaceous green in KOH.

M. staudtii Henn. var. staudtii differs by smaller basidiospores (20–)22.5–28.5 x 4.5–6.0 μm and smaller cheilocystidia (15–26 x 7.5–11 μm). Among African tropical species, M. zenkeri Henn. seems to be very close. It differs by violaceous tinged lamellae, the absence of pleurocystidia and smaller basidiospores ((15.5–)17.7–23.1 x 4.2–5.6 μm). Marasmius musisporus Desjardin & E. Horak from Papua New Guinea differs by a smaller (8–25 mm), greyish lilac pileus with yellow ridges, greyish lilac lamellae, a smaller stipe (50–70 mm long), narrower basidiospores (30–40 x 4.5–5.0 μm), smaller cheilocystidia (15–25 x 8–14 μm) and by the absence of pleurocystidia; M. purpureostriatus Hongo, collected in Japan and Papua New Guinea, has more distant lamellae (L = 10–13), smaller basidiospores (19–28(–32) x 4–6 μm), smaller cheilocystidia (15–37 x 10.0–15 μm), no pleurocystidia and well-developed broadly clavate caulocystidia (isotype ZT!).

In a note, J. Rammeloo writes that it is considered not edible by local people, and is known under the local name “muhungulanguba”.

 

55. Marasmius camerunensis Antonín & Mossebo

Antonín & Mossebo in Antonín, Mycotaxon 85: 113 (2003). Type: Cameroon, Littoral Province, near the village of Poola´a, ca. 5 km from Nkongsamba, 20 Aug. 1998, D.C. Mossebo M 196(1) (BRNM 670732, holotype; herb. Mossebo, isotype).

 

Selected descriptions and icons. Antonín, Mycotaxon 85: 113–114 (2003).

 

Pileus 40–70 mm broad, convex, deeply sulcate-striate, slightly rugulose except for a smooth area of 10–15 mm diam. at centre, whitish to ochraceous with a slightly umbonate brown-violaceous centre. Lamellae distant, L = 15, l = 0, adnexed, 8–10 mm broad, ventricose, intervenose, white, with concolorous entire edge. Stipe 40–70 x 4–6 mm, cylindrical, central, hollow, smooth, whitish and almost concolorous with lamellae at apex, reddish-brown towards base. Context concolorous with pileus, 2–5 mm broad in pileus centre. — Pl. 9

 

Basidiospores (21–)23–28(–30) x 5.5–6.5(–7.0) μm, E = 3.3–4.7, Q = 4.2, clavate, lacrimoid, clavate-cylindrical, thin-walled, hyaline, rarely uni- to triseptate. Basidia not found. Basidioles 13–35 x 8.5–10 μm, cylindrical, clavate, subfusoid. Cheilocystidia 10.0–25 x 8.5–11 μm, clavate, broadly clavate, rarely subcylindrical, thin-walled, hyaline. Pleurocystidia absent. Hyphae cylindrical, thin-walled, hyaline, 2.5–10.0 μm wide. Pileipellis a hymeniderm composed of 17–27 x 8.0–18 μm, clavate to broadly clavate, thin- to slightly thick-walled, hyaline to pale greyish-yellowish, smooth cells. Caulocystidia absent. Clamp-connections present in all tissues. – Fig. 59

 

Chemical reactions. All hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Single, lignicolous, on dead branches in a coffee plantation.

 

Distribution. Known only from one locality in Cameroon.

 

Revised specimens from tropical Africa.

Cameroon. Littoral Province, near the village of Poola´a, ca. 5 km from Nkongsamba, 20 Aug. 1998, D.C. Mossebo M 196(1) (BRNM 670732, holotype; herb. Mossebo, isotype).

 

Notes. Marasmius camerunensis is characterised by having rather large carpophores with a pale coloured non-striate pileus, broad lamellae, large basidiospores, and by the absence of pleurocystidia.

Among other African species with violaceous coloured or tinged pileus, M. bekolacongoli Beeli differs by a violaceous and yellow striped pileus, a longer stipe (80–150 x 2.5–6(–10) mm), smaller basidiospores (17.5–24.5(–26) x (3.8–)4.2–5.4 μm, Q = 5.1) and by growing on litter; M. violaceoides Antonín has entirely violaceous coloured or tinged carpophores and smaller basidiospores (15.5–22.3 x 3.3–5.0 μm); M. zenkeri Henn. has larger carpophores, a dark violaceous pileus, smaller basidiospores ((15.5–)17.7–23.1 x 4.2–5.6 μm) and larger cheilocystidia (14–50 x 10–15 μm). Marasmius caryotae (Berk.) Petch from Sri Lanka has a straw to greyish yellow pileus, a narrower stipe (40–80 x 1–2 mm), and shows fasciculate growth on fallen flowers of Caryota and on the ground (Pegler 1986, Petch 1948).

 

Sect. Sicci Singer

 

Marasmius sect. Sicci Singer, Mycologia 50:106 (1958).

Type species: Marasmius siccus (Schwein.) Fr.

 

Carpophores marasmioid or collybioid, small to medium-sized. Pileus usually membranaceous or thin-fleshed and mostly radially sulcate, white or pigmented, slightly pruinose or tomentose at least under lens. Lamellae distant to moderately crowded, thin, well-developed. Stipe non-insititious, narrow, cylindrical, usually tough, mostly with distinct basal mycelium. Rhizomorphs only rarely present.

 

Basidiospores moderately to very large (up to 45 μm long), clavate, fusoid or cylindrical-clavate; cheilocystidia present, usually in the form of broom-cells. Pleurocystidia present or absent. Pileipellis hymeniform, composed of broom-cells of the Siccus-type, with acute to obtuse projections, rarely composed of smooth cells and then setae present. Caulocystidia present or absent. Thick-walled setae may occur in pileipellis, hymenium, and stipitipellis. Hyphae of context dextrinoid (non-dextrinoid in subsect. Inaequales).

 

Notes. Section Sicci was proposed by Singer (1958a) for species with a pileipellis consisting of broom-cells of the Siccus-type and with dextrinoid trama hyphae. Earlier (Kühner 1933, 1936; Kühner & Romagnesi 1953), those species were included in sect. Globulares together with species with smooth elements in the pileipellis. It is clear that both sections (Globulares and Sicci) represent a uniform group; a division based on the presence or absence of broom-cells and smooth cells in pileipellis is quite artificial.

Singer (1964, 1965a) included Marasmius piperodorus Beeli to this section. However, the revision of the type specimen (BR 11501–55, holotype!) showed that it has not a hymeniform pileipellis of diverticulate hyphae and broom-cells, and all trama and context hyphae are non-dextrinoid. Therefore, this species belongs to the genus Marasmiellus.

 

 Key to tropical African species

 

1. Setae present on pileus, lamellae and/or stipe surface (ser. Spinulosi) ..................................................  11

1*. Setae absent ............................................................................................................................................  2

 

2. Stipe glabrous; caulocystidia absent (if present, then especially at apex and only in the form of broom-cells) ......  3

2*. Stipe entirely pruinose; caulocystidia present, cylindrical, clavate, subulate, thin- to thick-walled (ser. Atrorubentes) ....... 4

 

3. Pleurocystidia absent (ser. Leonini) .............................................  16

3*. Pleurocystidia present, mostly refractive (ser. Haematocephali) .............................  34

 

4. Carpophores entirely yellow; pleurocystidia present, 41–100 x 8.0–10.5 μm, fusoid, lageniform, subulate, rostrate; pileocystidia

present ................................................................  56. M. afrosulphureus

4*. Carpophores never entirely yellow; pleurocystidia mostly absent, if present then only up to 40 μm long, never subulate and rostrate; pileocystidia absent ................................  5

 

5.  Basidiospores larger than 11.5 μm ....................................................  6

5*. Basidiospores smaller than 12.5 μm ............................................. 7

 

6.  Pileus ochraceous to brownish; lamellae very crowded with concolorous edge; stipe white to pale alutaceous at apex, chestnut brown towards base; basidiospores 11.5–14 x 4.5–5.5 μm; pleurocystidia absent; caulocystidia 35–50 x 4.0–9.0 μm, cylindrical, narrowly clavate, sometimes irregular, simple, branched or with projections at apex, thick-walled ................................................. 57. M. xestocephalus

6*. Pileus brightly rusty ferrugineous to orange-ferrugineous, more fuscous at centre; lamellae moderately close with brown edge; stipe cinnamon-brown with paler apex; basidiospores (10–)13.5–18 x 3.5–4.5 μm; pleurocystidia present, 22–38 x 5–7 μm, fusoid to cylindrical-fusoid; caulocystidia 26–200 x 7.5–20 μm, subulate to lageniform, rostrate, (sub)acute, thin- to distinctly thick-walled (up to 1.5 μm) ..........................................................  59. M. atrorubens

 

7. Pileus red-brown, becoming fleshy pink with darker, brown centre; basidiospores very small, 4.5–6.5 x 2.7–3.5 μm; cheilocystidia irregular or lobate, smooth or with scattered apicular projections, never in the form of broom-cells; pileipellis consisting of both smooth and broom-cells ................. 61. M. buzungulo

7*. Pileus differently coloured, white, ochraceous to pale brown, reddish brown, orange brown; basidiospores always larger; cheilocystidia present, in the form of broom-cells, smooth-cells or a mixture of broom- and smooth cells ....................  8

 

8. Pileus white, sometimes pale ochraceous at centre, smooth; stipe often densely cespitose; basidiospores 6.0–8.0 x 3.0–3.2 μm ...............................  60. M. subarborescens

8*. Pileus always coloured, ochraceous to pale brown, reddish brown, orange brown; stipe sometimes in dense clusters, never densely cespitose; basidiospores either distinctly longer or broader ...................... 9

 

9. Pileus ochraceous to pale brown; basidiospores 8.0–9.5 x 3.7–4.2 μm; cheilocystidia lageniform, (sub)fusoid, irregular, mostly rostrate, never in the form of broom-cells; pileipellis composed of a mixture of broom- and smooth cells ................................... 58. M. xestocephaloides

9*. Pileus darker coloured, brownish orange, (reddish) brown, ochraceous-orange, brightly orange, yellow-orange; basidiospores either smaller (5.5–8.0 x 2.5–4.0 μm) or larger (7.0–12.7 x 3.0–4.6 μm); cheilocystidia in the form of broom-cells or both smooth and broom-cells; pileipellis only consisting of broom-cells ................................. 10

 

10. Pileus 13–35 mm broad, convex, distinctly radially rugulose around low umbo, brownish orange, ochraceous-orange to brightly orange at centre, paler, yellow orange or almost white towards margin; stipe pale cream above, through a pale brown zone up to rather dark (reddish) brown towards base; basidiospores 5.5–8.0 x 2.5–4.0 μm; cheilocystidia of both smooth and broom-cells; caulocystidia of one type: clavate, utriform, (sub)cylindrical, (sub)fusoid, lageniform, often irregular, simple or with lobate to diverticulate apex, never in the form of broom-cells ............  62. M. corrugatiformis

10*. Pileus 11 mm broad, conical, with prominent umbo, only slightly radially rugulose (especially at centre), dark (reddish) brown at centre, pale brown towards margin; stipe pale cream, with pale brown base; basidiospores 7.0–12.7 x 3.0–4.6 μm; cheilocystidia only in the form of broom-cells; caulocystidia of two types: cylindrical, sublageniform, subfusoid, subutriform, irregular or branched (especially at apex), and in the form of broom-cells (especially at centre) ............................. 63. M. katangensis

 

11. Basidiospores shorter than 10.5 μm ............................................................  12

11*. Basidiospores longer than 10 μm............................................................ 14

 

12. Pileus reddish brown; cheilocystidia clavate to utriform or ventricose, refractive, never in the form of broom-cells; pleurocystidia 25–45 x 8.0–10 μm, clavate, refractive; caulocystidia setoid ..................................  64. M. mengoënsis

12*. Pileus distinctly paler, white, yellowish, pale cinnamomeous brown or ochraceous; cheilocystidia always in the form of broom-cells, but sometimes poorly developed and together with smooth cells; pleurocystidia and caulosetae present or absent ....... 13

 

13. Cheilocystidia smooth or in the form of poorly developed broom-cells; basidia 13–18 μm long; pileipellis consisting of smooth cells; pileosetae present, caulosetae and hymenial setae absent .............................. 65. M. setiger

13*. Cheilocystidia in the form of well-developed broom-cells; basidia 20–27 μm long; pileipellis of broom-cells; pileo-, caulosetae and hymenial setae present ....................................................... 66. M. jalapensis

 

14. Cheilocystidia fusoid, (sub)cylindrical, lageniform, often rostrate, often moniliform, thin-walled, never in the form of broom-cells ....................................................... 67. M. pseudotorquescens

14*. Cheilocystidia always in the form of broom-cells ................................................... 15

 

15. Pileus umbrinous at centre, becoming chestnut brown towards margin; pleurocystidia 28–45 x 8.0–12 μm, clavate to subfusoid ................................................................................... 68. M. castaneovelutinus

15*. Pileus paler, yellowish brown, brown or red-brown; pleurocystidia 30–75 x 9.0–16 μm, versiform, (sub)fusoid, cylindrical, clavate, often rostrate ......................................................  69. M. fulvovelutinus

 

16.  Basidiospores shorter than (12–)13 μm ....................................................................... 17

16*. Basidiospores longer than (12–)13 μm .....................................................................  23

 

17. Basidiospores broader than 5.0 μm ........................................................................... 18

17*. Basidiospores narrower than 5.0 μm ..............................................................  19

 

18. Stipe long and wide (90–100 x 5–6 mm), distinctly broadened at base (9 mm), ochraceous yellow to yellowish brown; projections of pileipellis cells up to 15 μm long .................................. 70. M. episemus

18*. Stipe shorter and narrower (75 x 0.75 mm), cylindrical or only slightly broadened at base, yellowish white at apex, dark brown towards base; projections of pileipellis up to 9 μm long ..........................  71. M. ferruginacies

 

19. Pileus very small, (1–)2–4 mm broad, puniceous; stipe very thin, 0.2 mm wide; projections of pileipellis broom-cells numerous (c. 25–30) .......................................................................................... 72. M. leptus

19*. Pileus larger, at least 6 mm broad, differently coloured; stipe wider than 0.5 mm; projections of pileipellis broom-cells less numerous (up to ca. 25, only exceptionally up to 30) .................................  20

 

20. Pileus white, sometimes pale ochraceous at centre; basidiospores 6.0–8.0 x 3.0–3.2 μm; cheilocystidia inconspicuous, 8.0–15 x 4.0–9.0 μm, clavate, cylindrical-clavate, smooth or with 1–6 thin-walled, obtuse projections ............................................................................................................  60. M. subarborescens

20*. Pileus distinctly coloured; basidiospores longer than 7.5 μm and broader than 3.5 μm; cheilocystidia conspicuous, always in the form of well-developed broom-cells of the Siccus-type ...........................  21

 

21. Carpophores small (pileus 6–15 mm broad, stipe 5–10(–40) mm long); pileus ochraceous brown; stipe uniformly pale ochraceous; basidia and basidioles short (up to 24(–27) μm long) ........  73. M. ochropus

21*. Carpophores larger (pileus 5–30(–60) mm broad, stipe at least 30 mm long); pileus never entirely ochraceous brown (sometimes ochraceous brown only at margin); basidia and basidioles of the same size or longer .............  22

 

22. Pileus pale ochraceous buff, darkening to cinnamon at centre; lamellae intervenose, pale brown; stipe 0.5–3 mm wide; basidiospores 4.0–4.7(–5.5) μm broad ...............................................  74. M. bubalinus

22*. Pileus rusty brown; lamellae distant, pure white, not or only rarely intervenose (old carpophores); stipe 0.5–2 mm wide; basidiospores 3.5–4.2 μm broad ...................................................... 75. M. nodulocystis

 

23.  Basidiospores longer than (15–)16 μm .................................................................... 24

23*. Basidiospores 12–17 μm long ........................................................................ 31

 

24. Pileus white ................................................................  77.3. M. lilacinoalbus var. albus

24*. Pileus coloured ..................................................................................... 25

 

25. Basidiospores very large, 28–47 x 5.4–8.9 μm; basidia and basidioles very large, up to 65(–80) μm long ....  76. M. megistus

25*. Basidiospores distinctly smaller, up to 25 μm long and 6.0 μm wide; basidia and basidioles smaller, up to 48 μm long ..... 26

 

26.  Pileus distinctly radially striped ..............................................................................  27

26*. Pileus never striped ..............................................................................  29

 

27. Pileus with lilac or violet-brown colour .................................................  28

27*. Pileus without lilac or violet colours ........................................ 78. M. striaepileus

 

28. Striation dark violet-brown to lilac ..........................................  77.1. M. lilacinoalbus var. lilacinoalbus

28*. Striation lilac-purple, blood red or carmin red .................. 77.2. M. lilacinoalbus var. lilacinocarmineus

 

29. Pileus dull yellowish, orangish to rusty (purplish) brown; lamellae moderately distant, with darker edge; lamellulae usually absent; basidiospores 3.5–6.0 μm broad......................................... 79. M. sierraleonis

29*. Pileus differently coloured; lamellae distant; lamellulae present or absent; basidiospores 3.8–5.0(–5.5) μm broad .......  30

 

30. Pileus whitish cream at margin, ± ochraceous yellowish at centre; stipe distinctly yellow in upper part, dark brown towards base; basidiospores 16–18(–20) x 4.0–5.0 μm; basidioles up to 47 μm long .................... 80. M. luteostipitatus

30*. Pileus pinkish brown; stipe without distinct yellow colour in upper part; basidiospores 17.5–24 x 3.8–5.0(–5.5) μm; basidioles up to 38 μm long ................... 81. M. carcharus

 

31. Pileus white or pinkish, becoming pale buff or radially pale and white striate; lamellae narrow, often attenuated towards margin where they sometimes are reduced to veins or disappear ........................................... 82. M. haediniformis

31*. Pileus darker coloured, brown, orange-brown, reddish brown, orange-ochraceous .............................. 32

 

32. Pileus umbra or ± dark brown; lamellae dirty brown sometimes with pale brown edge; stipe 35–40 mm long; basidiospores 4.3–6.0 μm broad .......................................................................  83. M. macrolobieti

32*. Pileus brownish, orange-brown, orange-ochraceous; lamellae ochraceous cream or pale yellow, with concolorous edge; basidiospores 3.5–5.0 μm broad .............................................................................. 33

 

33. Pileus small, up to 5 mm broad, conical; lamellae moderately distant (L = 16–20) ....... 84. M. conicoparvus

33*. Pileus larger, 11–20 mm broad, broadly conical to convex, then conical to almost applanate; lamellae very distant (L = 10–16) .......................................  85. M. tanougouensis

 

34. Hyphae non-dextrinoid; pileus small (1.5–5 mm), with pale brown and dark ochraceous stripes; lamellae distant (L = 8–9), stipe filiform ..................................  101. M. beelianus

34*. Hyphae always dextrinoid; pileus larger; lamellae usually more close; stipe thicker ............................  35

 

35. Pileus purple, ruby, magenta, pastel red, purplish pink, purplish red, vinaceous red ....................  86. M. haematocephalus

35*. Pileus differently coloured, never purple, ruby, magenta, pastel red, purplish pink, purplish red, vinaceous red ............. 36

 

36. Stipe very long (20–150 mm); basidiospores very large, (17–)20–28 x 4.5–6.0 μm............... 87. M. longistipitatus

36*. Stipe shorter, up to 85 mm long; basidiospores shorter, up to 21.5(–23) μm long ...............................  37

 

37. Pileus white ..................................................................................  38

37*. Pileus always coloured .......................................................................  39

 

38. Pileus small to minute, up to 5 mm broad; lamellae very few (L = 8–9); stipe only 3–4 mm long, curved; pleurocystidia 26–35 x 9.0–11 μm large ....................................................... 88. M. robertsii

38*. Pileus 10–23 mm broad; lamellae more numerous (L = 10–14); stipe 20–45 mm long, straight; pleurocystidia 25–60 x 7.0–11 μm ................................. 89. M. haedinus

 

39. Pileus rusty-tawny or brown and pale tawny, with yellowish grey or orange stripes ...........................  40

39*. Pileus never striped ............................................................  41

 

40. Pileus brown with yellowish grey or orange stripes; lamellae distant (L = 16–18), with mostly coloured edge; stipe 15–85 x 0.5–2 mm; basidiospores 16.9–21.5(–23) x 3.5–5.4 μm ......  90. M. grandisetulosus

40*. Pileus rusty-tawny with paler tawny stripes; lamellae less numerous (L = 13–16), always with concolorous edge; stipe 15–30 x 0.5–1 mm; basidiospores 13.5–19 x 3.0–4.5 μm ..................  91. M. tenuisetulosus

 

41. Basidiospores 7.3–11 μm long ..............................................  42

41*. Basidiospores larger ............................................................  46

 

42. Basidiospores 7.3–9.2 x 3.6–4.6 μm; pileus brown, orange or brownish orange; caulocystidia in the form of broom-cells present ....................................... 99.3. M. confertus var. parvisporus

42*. Basidiospores 8.0–11 μm long; pileus grey-brown, orange-ferrugineous, cream, or yellow-ochraceous; caulocystidia in the form of broom-cells absent ...................................................................................  43

 

43. Pileus grey-brown; stipe orange to dark red towards base ..................................... 92. M. rubrostipitatus

43*. Pileus orange-ferrugineous, cream coloured or yellow-ochraceous; stipe never with orange or red tinges .................. 44

 

44.  Pileus 10–30 mm broad, orange-ferrugineous; stipe 25–60 x 1–3 mm, umbrinous to black with paler apex; basidiospores 8.0–10 x (2.5–)3.0–3.5(–4.0) μm; pleurocystidia 35–55 x 10–21 μm; pileipellis consisting of two types of broom-cells ........................................................................  93. M. spegazzinii

44*. Pileus up to 24 mm broad, yellow-ochraceous or cream coloured; stipe up to 40 mm long, pale brownish, cream or buff coloured at apex, up to brownish to sienna brown towards base; basidiospores up to 11 x 3.5–4.5 μm; pleurocystidia 32–59 x 7.0–17 μm; pileipellis consisting of one type of cell .................... 45

 

45. Pileus 14–24 mm broad, cream coloured; stipe up to 35 mm long, cream or buff coloured at apex, through golden or fulvous to sienna brown towards base; basidiospores (8.7–)9.0–11.5 x 3.5–4.2(–4.7) μm, fusoid, sublacrimoid; cheilocystidia in the form of broom-cells mixed with scattered smooth cells; pleurocystidia 32–59 x 8.0–17 µm ..... 94. M. cremeopileatus

45*. Pileus up to 10 mm broad; yellow-ochraceous; stipe up to 40 x 0.7 mm, pale brownish; basidiospores 8.0–11 x 3.5–4.5 μm, ellipsoid-fusoid, pip-shaped; cheilocystidia only in the form of broom-cells; pleurocystidia 37–57 x 7.0–12 μm ......................... 95. M. pallidopileatus

 

46. Pileus olive brown when young, then greyish yellow, blond to olive brown with yellowish brown margin; basidiospores 10.8–13.8(–14.5) x 3.8–5.4 μm; pleurocystidia 19–47 x 6.9–11(–13) μm; growing in close groups or cespitose ................................  96. M. elaeocephalus

46*. Pileus never with olivaceous tinge; basidiospores and pleurocystidia of different size; never growing in close groups or cespitose .................................................................  47

 

47. Basidiospores shorter than 14 μm and pleurocystidia narrow, 6.2–9.0 μm wide ................................  48

47*. Basidiospores longer than 13 μm or, if shorter, then pleurocystidia always wider than 9.0 μm ..........  49

 

48. Pileus campanulate, then broadly campanulate, deep yellow to yellowish orange; lamellae close, L = 18–21, with concolorous edge; stipe 30–65 x ± 1 mm; basidiospores 11.5–14 x 3.8–5.2 μm ..............................................  97. M. ferruginoides

48*. Pileus broadly conical to conical-convex, dark (reddish) brown, paler at margin; lamellae distant, L = 12, with dark brown edge; stipe 30–35 x up to 1 mm; basidiospores 10–12.5 x 3.0–4.3 μm .................................................  98. M. irangianus

 

49. Caulocystidia in the form of broom-cells present; basidiospores 11.5–15(–17) x 4.0–5.0(–6.0) μm; pileipellis consisting of one or two types of broom-cells ...........................................................  50

49*. Caulocystidia in the form of broom-cells absent; basidiospores 14–19(–21) μm long .........................  51

 

50. Pileus radially striate up to ½ of diameter; pleurocystidia 25–60(–75) x 10–16(–20) μm; pileipellis consisting of two types of broom-cells .....................................  99.1. M. confertus var. confertus

50*. Pileus only slightly striate at margin; pleurocystidia scattered, 25–45(–55) x 7.0–10 μm; pileipellis consisting of one type of broom-cell .........................................  99.2. M. confertus var. tenuicystidiatus

 

51.  Pileus striate at least on margin or up to ¾ of diameter, dark brown or chestnut black; lamellae close or subclose, L = 16–20; basidiospores 14–16(–17.5) x 3.8–4.5(–5.0) μm ............................ 100. M. strigipes

51*. Pileus brown, reddish brow, mahagony brown; lamellae more distant, L = 11–14 .... 101. M. bingaensis

 

 

Subsect. Siccini Singer

 

Subsect. Siccini Singer, Sydowia 18: 343 (1965).

 

Note. Trama hyphae always dextrinoid.

Series Atrorubentes

 

Ser. Atrorubentes Desjardin & E. Horak, Bibl. Mycol. 168: 27 (1997).

Ser. Actinopodes Singer p. p., Fl. Neotropica Monogr. 17: 236 (1976).

Type species: Marasmius atrorubens (Berk.) Berk. [Agaricus atrorubens Berk.]

 

Stipe pruinose or pubescent.

 

Pileipellis a hymeniderm composed of broom-cells of the Siccus-type. Caulocystidia present, cylindrical, clavate to attenuate, thin- or thick-walled. Pileo- and caulosetae and hymenial setae absent.

 

Note. The limit between ser. Atrorubentes and ser. Spinulosi is very narrow as for instance the type specimen of the type species, Agaricus atrorubens Berk., has both developed thin- and subulate thick-walled caulocystidia but typical setae are not developed. For details of delimitation of this, see Desjardin & Horak (1997).

 

Species descriptions

 

56. Marasmius afrosulphureus Courtec.

Courtecuisse, Doc. Mycol. 14(54–55): 90 (1984). Type: Kenya, Central Province, South Nyeri District, south side of Mt. Kenya, Castle Forest Station, 1 April 1968, D.N. Pegler K 321 (K(M) 92580, holotype). – Marasmius sulphureus Pegler, Kew Bull. Addit. Ser. 6: 186 (1977), non M. sulphureus A.E. Johnson, Bull. Minn. Acad. Nat. Sci. 1877: 337 (1878).

 

Selected descriptions and icons. Courtecuisse, Doc. Mycol. 14(54–55): 89–90 (1984); Pegler, Kew Bull. Addit. Ser. 6: 186 (1977) (as M. sulphureus).

 

Pileus 5–15 mm broad, convex, smooth, glabrescent, non-striate, uniformly sulphur yellow. Lamellae distant, L = 3–9, sometimes reduced to ridges, with occasional lamellulae, adnexed, concolorous with pileus. Stipe 5–10 x 1–2 mm, cylindrical, short, eccentric, curved, solid, pruinose, concolorous with pileus; arising from a cream coloured basal mycelium. Context white, thin. (According to Pegler 1977). — Pl. 10

 

Basidiospores 17–21 x 3.5–5.5 μm, E = 3.2–5.1, Q = 4.3, clavate, subfusoid, fusoid-clavate, sublacrimoid, thin-walled, smooth. Basidia 33–35 x 9.5–11 μm, 4-spored, clavate. Basidioles 18–35 x 5.0–11 μm, cylindrical, clavate, fusoid. Broom-cell cheilocystidia absent; lamella edge fertile, with scattered basidia mixed with ± clavate, (broadly) fusoid, subcylindrical, mostly irregular cystidioid cells mixed with scattered cystidia similar to pleurocystidia. Pleurocystidia numerous, 41–100 x 8.0–10.5 μm, fusoid, lageniform, subulate, rostrate, obtuse, thin-walled. Trama hyphae cylindrical to subinflated, ± thin-walled, smooth or minutely incrusted, up to 15 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 12–25 x 7.5–13 μm, clavate, (sub)cylindrical, thin- to slightly thick-walled, with digitate, slightly irregular, obtuse, slightly thick-walled, up to 15 x 3.0 μm projections. Pileocystidia numerous, 42–64 x 3.5–8.0 μm, narrowly lageniform, fusoid, slightly thick- to thin-walled, sometimes branched. Caulocystidia 30–300 x 2–6 μm, numerous, filiform, hyaline, thin-walled, sometimes branched. – Fig. 60

 

Chemical reactions. Trama, context and stipitipellis hyphae weakly to distinctly dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on fallen twigs on a forest floor.

 

Distribution. So far known only from Kenya.

 

Revised specimens from tropical Africa.

Kenya. Central Province, South Nyeri District, south side of Mt. Kenya, Castle Forest Station, 1 April 1968, D.N. Pegler K 321 (K(M) 92580, holotype).

 

Notes. Marasmius afrosulphureus has a unique position in this section by its entirely sulphur yellow carpophores, the absence of cheilocystidia in the form of broom-cells and the presence of pileocystidia and caulocystidia.

Pegler (1977) in original description of M. sulphureus mentioned slightly shorter and narrower basidiospores (11.5–18.2 x 3–4.5 μm), smaller basidia (21–27 x 6–8 μm) and smaller pleurocystidia (40–50 x 2–7 μm). Moreover, he did not mention the presence of cystidioid elements in the lamella edge. However, Courtecuisse (1984) mentioned the presence of the irregular, coralloid to diverticulate cheilocystidia.

Having entirely yellow carpophores with a short excentric stipe, well-developed pileo-, pleuro- and very long caulocystidia, M. afrosulphureus has a unique position among Marasmius species. Marasmius bellus Berk., known from Bolivia and Brazil, also has a yellow (“sunrise yellow”) pileus. However, its carpophores are larger (pileus 13–35 mm broad), with a central and larger stipe (35–54 x 1–1.2 mm), it has smaller basidiospores (8–12.7 x 3–4.8 μm) and lacks pleuro-, pileo- and caulocystidia (Singer 1976). Marasmius pseudoarachnoideus Dennis (= Amyloflagellula pseudoarachnoidea (Dennis) Singer), described from Trinidad and known also from Venezuela, is similar by the habit of its carpophores. However, it lacks cystidia, its carpophores are white to pale buff coloured and rhizomorphs are well-developed (Dennis 1951, 1970).

 

57. Marasmius xestocephalus Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 367 (1964). Type: Democratic Republic of Congo, Yangambi, J. Louis 14938 (BR 11532–84, holotype).

 

Selected descriptions and icons. Pegler, Kew Bull. Addit. Ser. 6: 199–200 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 367–369 (1964); Singer, Flore Icon. Champ. Congo 14: 272–273 & Pl. 45, fig. 3 (1965).

 

Pileus 10–18 mm broad, convex, obtusely umbonate, umbo sometimes in a central depression, smooth or slightly striate or sulcate at margin, always smooth at centre, glabrous, inflexed at margin when young, ochraceous to brownish (in alcohol between “burnt umber” and “kis kilim”, in herbarium “rust, sorolla” and “Alamo”, Maerz & Paul), often slightly darker at centre. Lamellae very crowded, l = 2–3, narrow, free or obtusely adnexed, not intervenose, pale alutaceous or pale ochraceous, with concolorous or paler, entire or slightly eroded edge. Stipe 12–28 x 0.7–1.2 mm, (almost) cylindrical, hollow, often curved, entirely slightly pubescent, glabrescent when old, white to pale alutaceous at apex, chestnut brown towards base, entirely brown when old; with rich fibrillose or strigose, white or brownish basal mycelium. Context white to whitish, thin. (According to Singer 1964, 1965a). — Pl. 10

 

Basidiospores 11.5–14 x 4.5–5.5 μm, E = 2.2–2.9, Q = 2.5, ± fusoid, thin-walled, hyaline, smooth. Basidia 27–30 x 8.0–10 μm, 4-spored, clavate. Basidioles 13–28 x 4.0–9.0 μm, cylindrical, clavate or fusoid. Cheilocystidia in the form of broom-cells of the Siccus-type, 12–17 x 6.0–9.0 μm, clavate or subcylindrical, thin-walled, with nodulose, thin- to slightly thick-walled projections. Pleurocystidia absent. Trama hyphae cylindrical, ± thin-walled, hyaline, up to 12 μm wide. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 9.0–20 x 6.0–9.0 μm, clavate to cylindrical, entirely thin-walled or thin-walled only at base and slightly thick-walled at apex, with 7–20 nodulose, digitate, obtuse to subacute, thin- to distinctly thick-walled projections, up to 8.0 x 1.0 μm (thin-walled) or 15 x 2.0 μm (thick-walled); thick-walled parts with ochraceous yellow walls in KOH. Stipitipellis a cutis composed of cylindrical, parallel, slightly thick-walled, up to 7.0 μm wide hyphae with ochraceous yellow walls in KOH. Caulocystidia 35–50 x 4.0–9.0 μm, cylindrical, narrowly clavate, sometimes irregular, simple, branched or with projections at apex, thick-walled, (sub)hyaline; mixed with scattered typical broom-cells at apex. Clamp-connections present in all tissues. – Fig. 61

 

Chemical reactions. Trama, context and stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on fallen twigs and lianas in a tropical forest in dry conditions (“terre ferme”).

 

Distribution. So far known only from the Democratic Republic of Congo and probably Ghana and Nigeria.

 

Revised specimens from tropical Africa.

Democratic Republic of Congo. Ipamu, H. Vanderyst 11077 (BR 11532–86, isotype); Yangambi, J. Louis 14938 (BR 11532–84, holotype).

Ghana. ? Cape Coast, near Jukwa, 10 May, A.C. Rose CC 6724 (K(M) 134628).

Nigeria. ? Cross River State, Calabar–Cameroon Road, 4 Aug. 1990, R.A. Nicholson 698 (K(M) 23103).

 

Notes. Marasmius xestocephalus is characterised by having an ochraceous to brownish pileus, moderately large, fusoid basidiospores, well-developed ± cylindrical to narrowly clavate, thick-walled, often (especially at apex) branched caulocystidia and by the absence of pleurocystidia.

In comparison with our observations, Singer (1964, 1965a) mentioned narrower basidiospores ((10–)11–14.5(–15.5) x (2.5–)3–4.2 μm) and larger pileipellis cells (4.8–34 x 7–25 μm).

The closest species seems to be Marasmius ochroleucus Desjardin & E. Horak, described from New Caledonia. It differs in having a paler, off-white, pale ochraceous or pale orange-ochraceous pileus, an entirely pale yellowish brown stipe, smaller basidiospores (9–11 x 3.5–4 μm) and its caulocystidia being only sometimes apically thick-walled (Desjardin & Horak 1997).

 

58. Marasmius xestocephaloides Antonín

Antonín, Mycotaxon 89(2): 420 (2004). Type: Kenya, Central Province, Nairobi District, Thika, Thika Falls, 16 March 1968, D.N. Pegler 94 (K(M) 116841, holotype).

 

Misapplication: Marasmius xestocephalus Singer s. Pegler (1977).

 

Selected descriptions and icons. Antonín, Mycotaxon 89(2): 420-422. 2004; Pegler, Kew Bull. Addit. Ser. 6: 199–200. 1977.

 

Pileus 10–30 mm broad, convex, subumbonate, smooth, glabrous, ochraceous to light brown, darker at centre. Lamellae very crowded, l = 3, free to adnexed, narrow (up to 1 mm), pale alutaceous to ochraceous. Stipe 15–40 x 1–1.5 mm, cylindrical, often curved, hollow, finely pruinose, whitish above, chestnut brown below; with whitish strigose mycelium at base. Context thin, whitish. (According to Pegler 1977). — Pl. 10

 

Basidiospores 8.0–9.5 x 3.7–4.2 μm, E = 2.0–2.6, Q = 2.2, ellipsoid-fusoid, fusoid, thin-walled, smooth, hyaline. Basidia 20–24 x 5.0–7.5 μm, 4-spored, clavate. Basidioles 15–21 x 4.0–7.0 μm, clavate, cylindrical, fusoid. Cheilocystidia 18–35 x 5.0–8.5 μm, lageniform, (sub)fusoid, mostly rostrate, irregular, thin-walled, hyaline. Pleurocystidia absent. Trama hyphae cylindrical to subinflated, thin-walled, hyaline, up to 15 μm wide, mixed with slightly thick-walled, up to 8.0 μm wide ones. Pileipellis a hymeniderm composed of (1) broom-cells of the Siccus-type, sometimes transient to the Rotalis-type, sometimes (almost) smooth, 12–22 x 7.0–16 μm, clavate, pyriform, cylindrical or vesiculose, thin- to slightly thick-walled, with up to 10 x 1.5 μm, ± slightly thick-walled, obtuse and smooth projections, and (2) up to 65 x 8.0 μm, distinctly thick-walled broom-cells transient to setoid cells (however, true setae absent!); subpileipellis composed of ± globose cells. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled, up to 8.0 μm wide hyphae, with subhyaline to pale yellowish walls in KOH. Caulocystidia numerous, 25–40 x 7.0–11 μm, cylindrical, subfusoid, subclavate, ± thick-walled, with subhyaline to pale yellowish walls in KOH. Clamp-connections present in all tissues. – Fig. 62

 

Chemical reactions. Trama, context and stipitipellis hyphae dextrinoid, other structures and basidiospores non-dextrinoid.

 

Ecology. Saprophytic, growing in ± dense groups on dead wood and forest debris (e.g. Sysygium sp.).

 

Distribution. So far known only from Kenya, Uganda and probably from Zambia.

 

Revised specimens from tropical Africa.

Kenya. Central Province, Nairobi District, Thika, Thika Falls, 16 March 1968, D.N. Pegler 94 (K(M) 116841, holotype as M. xestocephalus).

Uganda. Buganda Province, Mengo District, Mawakota County, Mpanga Reserve Forest, 8 June 1968, D.N. Pegler U 1330 (K(M) 8874, as M. xestocephalus).

Zambia. ? Chowo Forest, 12 Dec. 1981, J. Rammeloo 7843 (BR 12027–96).

 

Notes. Marasmius xestocephaloides is characterised by having an ochraceous to light brown pileus, very crowded lamellae, a chestnut brown stipe at base, small basidiospores, short basidia and basidioles, cheilocystidia are lageniform, (sub) fusoid and mostly rostrate and irregular, never in the form of broom-cells, a pileipellis composed of both ± thin-walled broom-cells and (almost) smooth cells, mixed with distinctly thick-walled broom-cells transient to setoid cells and in having cylindrical, subfusoid or subclavate, ± thick-walled caulocystidia. Pleurocystidia as well as true setae are not developed.

Collection J. Rammeloo 7843 from Zambia is very similar both macroscopically (± same colour of carpophores with very close narrow lamellae and growth in dense groups) and microscopically (pileipellis a mixture of broom- and smooth cells, similar caulocystidia). However, its cheilocystidia are in the form of broom-cells mixed with smooth, regular, lobate to subcoralloid cells. Therefore, it is included with a question-mark here.

Pegler (1977) mentioned distinctly larger basidiospores (11–14 x 3.2–4.5 μm), cheilocystidia in the form of broom-cells and did not mention the presence of thick-walled setoid cells in the pileipellis. Maybe he studied a mixed collection.

Marasmius xestocephalus Singer represents a very closely related species. It differs in having smaller carpophores, larger basidiospores (11.5–14 x 4.5–5.5 μm), longer basidia and basidioles (27–30 x 8.0–10 μm), cheilocystidia in the form of broom-cells and different pileipellis broom-cells. Marasmius subarborescens Singer has a white pileus, different cheilocystidia and even smaller basidiospores (6.0–8.0 x 3.0–3.2 μm).

The absence of cheilocystidia in the form of broom-cells represents a unique character in this series. Only Marasmius heterocheilus Singer, known from Bolivia, is described as having mostly clavate and simple, rarely cylindrical or clavate cheilocystidia with one to four apical projections, which are often irregularly contorted. However, it differs in having a cinnamomeous coloured, up to 50 mm broad pileus, a longer and more robust stipe (up to 80 x 4 mm), smaller basidiospores (6–6.3 x 3.5–4 μm), and differently shaped caulocystidia which have the same form as the cheilocystidia (Singer 1976).

 

59. Marasmius atrorubens (Berk.) Mont.

Montagne, Ann. Sci. Nat., Bot., sér. 4, 1: 118 (1854). Agaricus atrorubens Berk., Journ. Bot. 1: 138 (1842). Type: Surinam, Hostmann 297 (K(M) 99653, ex herb. Hooker, holotype). – Marasmius castaneus Mont., Ann. Sci. Nat., Bot., sér. 4, 1: 109 (1854). – Marasmius portoricensis Murrill, N. Amer. Fl. 9: 262 (1915).

 

Misapplication: Marasmius actinopus Mont., Ann. Sci. Nat., Bot., sér. 4, 1: 112 (1854) s. Singer (1964, 1965a).

 

Selected descriptions and icons. Montagne, Ann. Sci. Natur., Bot., sér. 4, 1: 109 (1854) (as M. castaneus); Mossebo & Antonín, Czech Mycol. 56(1-2): 86–90 & Pl. 1 (2004); Pegler, Kew Bull. Addit. Ser. 6: 196–197 (1977); Pegler, Kew. Bull. Addit. Ser. 9: 215–216 (1983); Singer, Bull. Jard. Bot. Etat Brux. 34: 377–378 (1964) (as M. actinopus); Singer, Flore Icon. Champ. Congo, fasc. 14: 275–276 (1965) (as M. actinopus).

 

Pileus (5–)10–17 mm broad, broadly conical with pronounced central obtuse-conical umbo and slightly involute margin, then (almost) applanate with small central obtuse umbo and slightly inflexed to straight margin, hygrophanous, slightly translucently striate, finely tomentose, pruinose, sometimes finely rugulose at centre, slightly striate at margin when old, not striate in young carpophores, entirely orangish brown (7C–D8) when young, then darker orangish brown (7D–E8, 8–9D8) at centre, paler (6–7C7 to 8C8) towards margin. Lamellae rather close, L = 14–17(–20), l = 2–4, emarginate, adnexed with a small tooth, rather narrow (c. 1.5 mm), neither intervenose nor branched, pale cream (± 4A2 to 5A2–3), with concolorous, entire, finely pubescent edge. Stipe (15–)25–40 x 0.5–1 mm, cylindrical, slightly broadened above, rarely slightly broadened at base, non-insititious, straight or slightly flexuose, entirely strigose-hairy, whitish at apex, reddish (orangish) brown (7–8D–E7–8 to 9D–E8) towards base; with large (up to 5 mm) basal mycelium of strigose, adpressed to erect, reddish brown hairs. Context almost absent in pileus, concolorous with surface, hollow in stipe, with fungoid smell and mild taste— Pl. 10

 

Basidiospores (10–)11.5–14(–18) x 3.6–5.0 μm, E = 2.5–3.5, Q = 2.5–3.1, clavate to (clavate-)fusoid, thin-walled, smooth, hyaline. Basidia 19–24 x 5.0–8.0 μm, 4-spored, clavate. Basidioles 11–30 x 4.0–11 μm, clavate, subcylindrical or subfusoid. Cheilocystidia in the form of broom-cells, 8.0–28 x 5.5–10 μm, clavate to cylindrical, ± hyaline, with thin or slightly thickened walls. Pleurocystidia 22–38 x 5.0–7.0 μm, fusoid to cylindrical-fusoid, often with mucronate apex, refractive, hyaline to pale yellowish, mostly thin-walled. Trama hyphae cylindrical, ± thin-walled, hyaline, up to 12 μm wide. Pileipellis a hymeniderm composed of (1) broom-cells of the Siccus-type, 8.0–23 x 4.0–10 μm, clavate to cylindrical-clavate, slightly thick-walled, less frequently thin-walled, hyaline or pale ochraceous yellow below, ferrugineous above, with (10–)15–30(–35) subacute, slightly thick-walled, ± nodulose, up to 9.0 x 1.5 μm projections, mixed with (2) 25–55 x 8.0–20 μm, setoid broom-cells with up to 100 μm long projections; walls of both types ochraceous ferrugineous or yellow-brown in KOH. Stipitipellis a cutis consisting of cylindrical, parallel, slightly thick-walled (up to 1.0 μm), up to 5.0 μm wide hyphae with ochraceous yellow walls in KOH. Caulocystidia numerous, 20–240 x 7.5–20 μm, subulate to lageniform, rostrate, (sub)acute, thin- to thick-walled (walls up to 1.5 μm), hyaline to pale ochraceous yellow, simple. Clamp-connections present in all tissues. – Fig. 63

 

Chemical reactions. Trama, context and stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing on dead leaves of Macrolobium dewevrei and other dicotyledons in a tropical forest.

 

Distribution. It seems to be widely distributed in tropical Africa. So far, it has been found in Benin, Cameroon, Democratic Republic of Congo, Tanzania, Uganda and probably also Burundi. It also occurs in South America.

 

Revised specimens from tropical Africa.

Benin. Atacora Province, Kota, 27 Aug. 1997, V. Antonín B97.115 (BR 101160–86); Ibid., 23 Aug. 1997, V. Antonín B97.91 (BR 101137–63); Ibid., 26 Aug. 1997, V. Antonín B97.104 (BR 101149–75); Borgou Province, Wari Maro, 22 Aug. 1997, V. Antonín B97.88 (BR 101134–60).

Burundi. ? Bururi Province, Kigwena, 20 Febr. 1979, J. Rammeloo 6680 (BR 11942–11).

Cameroon. Yaoundé, Mt. Eloundem, 30 March 2001, V. Antonín Cm 01.09 (BRNM 666058); South West Province, Korup National Park, trail from Rengo Camp to Ekunde–Kunde, 9 April 1997, P.J. Roberts K 996 (K(M) 92734, BRNM 686377); Ibid., P.J. Roberts K 995 (K(M) 91508, BRNM 686383); Ibid., transect P, 25 April 1996, P.J. Roberts K 120 (K(M) 92739, BRNM 686382); Ibid., Ekunde–Kunde, path to Ysuky, 26 April 1996, P. Slangen 127 (K).

Democratic Republic of Congo. 20 km NE of Yambao, 10 June 1939, J. Louis 15241 (BR 11373–24); Ibid., J. Louis 15216 (BR 11372–23);  Lemfu, H. Vanderyst 699 (? BR); 7 km from Yangambi, 15 Sept. 1957, B. Fassi 1109 (BR 11542–96); Tshopo Province, Kisangani, Arboretum at the University Campus, 24 April 1984, B. Buyck 1553 (BR 11743–06); Ibid., 24 April 1984, B. Buyck 1554 (BR 11742–05).

Tanzania. Dar es Salaam, Botany Department, 2 May 1976, C.M.R. Hennessy 196 (K(M) 116833); East Usambara Mts., Tanga Province, Lushoto District, Amani, 15 April 1968, D.N. Pegler T 469 (K(M) 134262).

Uganda. Buganda Province, Mengo District, 6 km N of Entebbe, Zika Forest, 12 June 1968, D.N. Pegler 1449 (K(M) 116828).

 

Revised specimens from other regions.

Surinam. Hostmann 297 (K(M) 99653, ex herb. Hooker, holotype).

 

Notes. Marasmius atrorubens is characterised by having an often papillate orangish brown (when young), then darker orangish brown pileus at centre, paler towards margin, rather close lamellae, a reddish (orangish) brown stipe, moderately large basidiospores, well-developed mucronate short pleurocystidia and numerous ± slightly thick-walled caulocystidia.

Having well-developed pleurocystidia, the type specimen ± agrees with the taxon M. atrorubens var. cystidifer Singer (Singer 1976). Pegler (1977, 1983) mentioned smaller basidiospores (10–15 x 2.5–3.8 μm) and the absence of pleurocystidia. Dennis (1951) found setoid elements in the pileipellis and proposed the new var. setosus Dennis.

Marasmius nummularius Berk. & Broome, found in Sri Lanka and Indonesia, differs in having concolorous lamellar edges, smaller basidiospores: (11–)12–15 x (3–)3.5–5 μm (Desjardin & al. 2000) or 10–12 x 3–3.5 μm (Pegler 1986), no pleurocystidia and well-developed caulosetae. Moreover, Desjardin & al. (2000) mentioned two types of cheilocystidia, Pegler (1986) only one. Marasmius glaucopus (Pat.) Sacc. & D. Sacc., described from Guadeloupe, has a dark purplish brown pileus, dark purple lamellae and smaller basidiospores (8.3–9.3 x 3.8–5 μm) (Pegler 1983; Singer 1976).

 

60. Marasmius subarborescens Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 364 (1964). Type: Democratic Republic of Congo, Eala, Feb. 1923, M. Goossens–Fontana 113 (BR 11511–65, holotype).

 

Selected descriptions and icons. Pegler, Kew Bull. Addit. Ser. 6: 196 (1977); Singer, Bull. Jard. Bot. Etat Brux. 34: 364–365 (1964); Singer, Flore Icon. Champ. Congo 14: 271–272 & Pl. 46, fig. 8 (1965).

 

Pileus 8–35 mm broad, hemispherical or campanulate, then campanulate-convex or plano-convex, margin involute when young and slightly reflexed when old, sometimes slightly depressed at centre, smooth, glabrous, white, sometimes pale ochraceous at centre. Lamellae very close to close, with numerous lamellulae, adnate to almost free, not intervenose, narrow. Stipe 23–60 x 0.5–1.5 mm, cylindrical to subcylindrical, often twisted, sometimes flexuose, glabrous-subpruinose, smooth, whitish or cream at apex, ochraceous brown or chestnut towards base, in basal part densely cespitose; with white basal mycelium. Context white, concolorous with surface in stipe base. (According to Pegler 1977, Singer 1964, 1965a). — Pl. 10

 

Basidiospores 6.0–8.0(–9.0) x 2.7–3.2 μm, E = 2.0–3.3, Q = 2.5, ellipsoid, subfusoid, sublacrimoid, thin-walled, smooth. Basidia e.g. 21.5 x 6.9 μm, 4-, rarely 2-spored, clavate. Basidioles 12–27 x 5.5–7.0 μm, clavate, cylindrical, subfusoid. Cheilocystidia inconspicuous, 8.0–15 x 4.0–9.0 μm, clavate, cylindrical-clavate, smooth or with 1–6 thin-walled, obtuse projections; sometimes scattered or absent(?). Pleurocystidia absent. Trama hyphae cylindrical or subinflated, ± thin-walled, (sub)hyaline, up to 15 μm wide, mixed with narrower (up to 5.0 μm), thick-walled ones. Pileipellis a hymeniderm composed of broom-cells of the Siccus-type, 14–15.5 x 6.9–8.5 μm, clavate to subcylindrical, thin-walled at base and slightly thick-walled above, with 10–20(–25) digitate, obtuse to subacute, slightly thick-walled, slightly nodulose, up to 6.2 x 1.0 μm projections; thick-walled parts with yellow-brown walls in KOH; mixed with less frequent smooth, ± clavate cells. Stipitipellis a cutis composed of cylindrical, parallel, slightly thick-walled, up to 6.0 μm wide hyphae, with yellow-brown walls in KOH. Caulocystidia numerous, especially towards apex, sometimes ± scattered and less distinct, 16–35 x 7.0–10 μm, adpressed to erect, clavate, subcylindrical, fusoid, ± thin-walled, sometimes mixed with scattered broom-cells; walls pale dirty yellow or subhyaline in KOH. Clamp-connections present in all tissues. – Fig. 64

 

Chemical reactions. Trama, context and stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, growing in dense groups, less frequently single, on decaying wood and litter in various types of forest and in plantations.

 

Distribution. Hitherto found in Angola, Cameroon, Democratic Republic of Congo, Ghana, Ivory Coast and Uganda.

 

Revised specimens from tropical Africa.

Angola. Golungo Alto, March 1855, Welwitsch 262 (K(M) 134445); Ibid., Apr. 1856, Welwitsch 266 (K(M) 134444).

Cameroon. Sud Province, Somalomo, Dja Biosphere Reserve, 12 April 2001, D.C. Mossebo 192D (herb. Mossebo).

Democratic Republic of Congo. Eala, Feb. 1923, M. Goossens–Fontana 113 (BR 11511–65, holotype); Ibid., M. Goossens–Fontana 11 (BR 11512–66, isotype).

Ghana. Gold Coast, 6 June 1915, R.H. Banting 15 (K(M) 134437).

Ivory Coast. Forêt de Tai, 9 Jan. 1976, L. Aké Assi 440 (K(M) 134438); Forêt de la Besso, 31 March 1975, L. Aké Assi 321 (K(M) 133805, as M. arborescens).

Uganda. Makerere College, Mpanga 69, 9 April 1964, E.A. Calder 34 (K(M) 133806, as M. corrugatiformis); Buganda Province, Mengo District, Mawakota County, Mpanga Research Forest, 8 June 1968, D.N. Pegler U 1324 (K(M) 134443); Ibid., D.N. Pegler U 1458 (K(M) 134440); Buganda Province, Mengo District, north of Entebbe, Zika Forest, 12 June 1968, D.N. Pegler U 1446 (K(M) 134439); Western Province, Bunyoro District, near Masindi, Budongo Forest, 15 June 1968, D.N. Pegler U 1468 (K(M) 134441); Ibid., 16 June 1968, D.N. Pegler U 1512 (K(M) 134442).

 

Notes. Marasmius subarborescens is characterised by having a white, sometimes at centre pale ochraceous pileus, very close to close lamellae, an often densely cespitose growth, small basidiospores, small cheilocystidia and well-developed caulocystidia. This combination of characters is quite unique in sect. Sicci. Caulocystidia are sometimes scattered and less distinct and may be overlooked; therefore, this species is also keyed out as having no caulocystidia.

Cheilocystidia were not found in the type collections because of the very bad condition of holotype and isotype specimens. Even Singer (1964) did not mention the presence of cheilocystidia. Pegler (1977) mentioned smaller basidia (12–16 x 4–5 μm) and smaller pileipellis cells (7–12 x 6–8.5 μm), and Singer (1964, 1965a) also slightly smaller basidia (11.7–16 x 3.5–4.5 μm) and pileipellis cells (5.5–10.5 x 4.5–7 μm). In addition, none of them mentioned the presence of caulocystidia.

 

61. Marasmius buzungulo Singer

Singer, Bull. Jard. Bot. Etat Brux. 34: 371 (1964). Type: Democratic Republic of Congo, vicinity of Kinshasa (as Léopoldville), Nov. 1947, L. Dubois 1501 (BR 11417–68, holotype).

 

Selected descriptions and icons. Singer, Bull. Jard. Bot. Etat Brux. 34: 371–373 (1964); Singer, Flore Icon. Champ. Congo 14: 273–274 (1965).

 

Pileus 30–40 mm broad, campanulate, then expanded, umbonate, scrobiculate-rugose at margin, red-brown, becoming fleshy pink, with darker, brown centre, sometimes cracked in old specimens, sometimes with whitish stains. Lamellae crowded, with many lamellulae, obtusely adnexed, then slightly decurrent, narrow to rather large, white, with entire, concolorous edge. Stipe 40–70 x 2–4 mm, cylindrical, hollow, often slightly twisted, sometimes curved, finely pruinose, white; with well-developed whitish basal mycelium. Context white, with mouldy smell and without taste but slightly astringent after-taste. (According to Singer 1964, 1965a). — Pl. 10

 

Basidiospores 4.5–6.5 x 2.7–3.5 μm, ellipsoid, cylindrical-ellipsoid, thin-walled, hyaline. Basidia 16 x 6.0 μm (one found), 4-spored, clavate. Basidioles 13–22 x 4.0–8.0 μm, cylindrical, clavate, subfusoid. Cheilocystidia 13–22 x 5.0–10 μm, ± clavate, often irregular, smooth, lobate or with apical projections, hyaline, thin-walled. Pleurocystidia absent. Trama hyphae cylindrical, ± thin-walled, hyaline, up to 10 μm wide. Pileipellis a hymeniderm composed of (1) smooth, (14–)18–28 x 8.0–13 μm, clavate, (sub)pyriform, subcylindrical, thin- to slightly thick-walled cells and (2) broom-cells of the Siccus-type, 17–36 x 8.0–15 μm, clavate to subcylindrical, slightly to distinctly thick-walled, sometimes irregular, with 3–10, obtuse, thick-walled, up to 10 x 3.0 μm projections; thick-walled parts with ochraceous yellow walls in KOH. Stipitipellis a cutis composed of cylindrical, parallel, slightly thick-walled, up to 5.0 μm wide hyphae, with ochraceous yellow walls in KOH. Caulocystidia scattered, 17–30 x 6.0–10 μm, adpressed to erect, cylindrical to clavate, thick-walled, simple or often with 2–5 apical projections, with ochraceous walls in KOH. Clamp-connections present in all tissues. – Fig. 65

 

Chemical reactions. Trama, context and stipitipellis hyphae dextrinoid, other structures non-dextrinoid.

 

Ecology. Saprophytic, in groups of 3 or 4 carpophores, growing on dead leaves and on detritus in a savannah-woodland.