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

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).

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.

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)

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).

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).