A revision of the genus Inocybe in Europe. I. Subgenus Inosperma and the smooth-spored species of subgenus Inocybe Th.W. Kuyper Rijksherbarium, Leiden Date of publication: 12 December 1986 BUT WHY THY ODOR MATCHETH NOT THY SHOW THE SOIL IS THIS, THAT THOU DOST COMMON GROW Shakespeare, Sonnet 69 The editors of the journal Persoonia gratefully acknowledge a substantial contribution by the 'Rijksherbarium Foundation Dr. E. Kits van Waveren' in the cost of printing of this Supplement Volume of Persoonia, thus making the publication of this work by the Rijksherbarium financially possible. © 1986 Rijksherbarium Leiden ISBN 90-71236-02-1 ISSN 0031-5850 Contents Introduction Acknowledgements A. 1 General part Chapter I. Material and methods 1 Chapter II. Notes on characters 2 — — — I. Habitat. — 2. Pileus. 3. Lamellae. 4. Stipe. 5. Context. — 6. Smell. 7. Velum. — 8. Ontogeny. — 9. Spores. — 10. Basidia. II. Clamp-connections. — 12. Cystidia. — 13. Pileipellis. 14. Chemistry. Chapter III. Ecology and distribution 13 1. Ecology. — 2. Distribution. — 3. Phenology. Chapter IV. Evolution of Inocybe 15 1. Affinities of with other — 2. Cladistic of Inocybe genera. analysis Inocybe. — 3. Infrageneric taxonomy of Inocybe. — 4. Species concept and speciation in Inocybe. References 26 B. Special part 29 Inocybe 29 Key to the subgenera 30 Synopsis 30 Synoptical key 32 Subgenus Inosperma 34 Section Cervicolores 34 Section Rimosae 43 Subgenus Inocybe 69 'Supersection'Cortinatae (species 17-69) 75 'Supersection' Marginatae (species 70-93) 180 Excluded species and nomina dubia 221 Index 240 Introduction Fries (1821: 11) established Agaricus series Derminus tribus Inocybe, which was later elevated by him to generic rank (Fries, 1863: 346). Originally Fries based his of circumscription Inocybe solely on macroscopical characters; when it was raised to status Fries added that the of generic spores all Inocybe species were seemingly rough (’sporae scabrae videntur omnibus Inocybis communes’). Almost certainly this based was not on original observation, as Fries considered the use of the microscope unnecessary, but the result of an uncritical appraisal of observations by Berkeley (1860). The relevance of Fries’s statement regarding the typification of is discussed on 29. Inocybe p. the is Although genus Inocybe easily recognisable by macroscopical characters, from the fact that the has the judging generic concept scarcely changed to present day, the delimitation of species is considerably more difficult. The number of species increased from 1821 continually onwards. Massee (1904) published a world mono- of the but his work and it graph genus unfortunately lacks precision is moreover outdated. Important regional work on the genus has been carried out by Heim (1931) and Kühner (in Kühner & Romagnesi, 1953) in France; Alessio (1980) in Italy; Enderle & Stangl (1981) and Stangl & Enderle (1983) in Germany; Kauffman (1924), Stuntz (1947, 1954), and Grund & Stuntz (1968, 1970, 1975, 1977, 1980, 1981, 1983, 1984) in North America; and Horak (1978, 1979, 1980, 1981) in Australia and Asia. A treatment of the Inocybe species occurring in the Netherlands has been given by Boedijn (1925), but this work has now become obsolete. Important contributions to the knowledge of the Netherlands's species have been made by Huijsman but the results of his studies have for the greater part not been published. Compilatory keys to all European species of Inocybe have been provided by Bataille (1910), Alessio (1980), Enderle & Stangl (1981), Stangl & Enderle (1983), and Moser (1983). A real disadvantage of compilatory keys, however, is that they result in a rather of It large overestimation the actual number of species. was therefore felt that of a critical revision the European species was desirable. However, due to the large number of species and the limited amount of time available for this study, I had confine of the in to myself to a study smooth-spored species occurring north- wester At later also excluded Europe. _a stage subgenus Mallocybe (see p. 22) was from this studv. On the other hand I tried to include revision of all a Inocybe- types from and also studied from described Europe, I several types North America, as the flora of both alike Inocybe continents appeared to be more than had hitherto been suspected (cf. Lange, 1934). ACKNOWLEDGEMENTS I feel much indebted to Dr C. Bas for his work encouragement during my on Inocybe; to Mr H. S. C. Huijsman (Beilen) and Mr J. Stangl (Augsburg) for their constant cooperation; to the directors and curators of the following herbaria BERN, BP, BRA, BRNO, C, CUP, E, FH, G, H, IB, K, KR, LE, M, MARS, MICH, MPU, NYS, O, PC, PRC, PRM, S, WBS, and WTU from which important collections were obtained on loan; to Mr M. Bon (Saint Valery-sur-Somme), Mr R. Courtecuisse (Aubers), Mr C. Furrer-Ziogas (Basel), Mr P. Reumaux (Paris), and Mr H. Romagnesi of from their (Paris) for the loan type specimens herbaria; to Prof. Dr C. Kalkman, Prof. Dr R. Kiihner (Lyon), and Drs E.C. Vellinga for constructive criticism of this manuscript; to Dr D. A. Reid (Kew) for linguistic improvement of the text of the all those well introductory chapters; to mycologists, amateurs as as pro- who with fresh collections and fessionals, supplied me many interesting well-annotated exsiccates; to Ruth van Crevel and Jan van Os for preparing the illustrations for printing; to the Netherlands Organization for the Advancement of Pure Research for financial in France in 1983. (Z.W.O.) support during my stay A. GENERAL PART CHAPTER I MATERIAL AND METHODS Most of the dealt with herein have been studied species by me in fresh as well in dried characters therefore as condition. Macroscopical are for the most part based on personal observations, supplemented by descriptions provided by other mycol- ogists. It has been explicitly indicated when descriptions have been taken from the literature. The colours of fresh specimens are compared with Munsell Soil Colour Charts, Baltimore, or with Kornerup & Wanscher, MethuenHandbook of Colour. examinations have Microscopical always been made in a 10% NH OH-solution. 4 with old in KOH-solution Only very specimens a pretreatment a 5% was sometimes necessary. Although both solutions yield a comparable colour of the cystidial wall in thick-walled cystidia (see p. 10), the use of the latter solution is to be discouraged as the spores tend to swell strongly, thereby leading to unrepresentative values. characters have been measured under oil Microscopical an immersion objective. Values of have been rounded to the 0.5 spores nearest /am, those of cystidia and basidia to the nearest 1 /am. The have been made with the aid of drawings a camera lucida. The magnifications of the figures are: spores x 1500, cystidia x 1000. The calcium oxalate-crystals at the have but their dimensions influenced cystidial apex been drawn, frequency and are the by age and mode of preservation of the material. Drawings of these could therefore well be atypical and lead to false conclusions. The abbreviations of the herbaria follow Holmgren & al. (1981). Unless otherwise indicated, collections mentioned are deposited at L. 1 CHAPTER II NOTES ON CHARACTERS 1. Habit the is The habit of basidiocarps in most species tricholomatoid or collybioid, Differences in habit rather difficult exceptionally mycenoid. are, however, to express and not sharply distinct and therefore of only limited value for specific discrimination. of defined In some cases an Index Slenderness may be useful, as l 2 IS " d. D (Heinemann, 1981), in which 1 is length of stipe, d is diameter of stipe and D is diameter of pileus. This index is often useful for separating the two varieties of I. phaeocomis and of I. splendens. Only mature specimens should be used in calculating this index. 2. Pileus Size and shape of the pileus are rather variable too and can therefore only rarely be used as a taxonomic character. is for a Shape large part age-dependent, young specimens generally being more or less conical or campanulate, and mature specimens being convex, plano-convex to applanate. An umbo is usually present, but is often This character lacking when the velipellis is well-developed (see p. 15). must therefore be used with caution. The colour shade of great is usually some yellow, brown other The or red-brown, tinges being diagnostically more important. pileus of only three species, viz. I. impexa, I. hygrophana, and I. ionochlora, is reported to be hygrophanous. The surface of the pileus is far more variable than the generic name (Inocybe 'fibrillose means pileus') implies and is taxonomically rather important, although Surface texture only at specific level. ranges from smooth to radially fibrillose, squamulose or squarrose. However, a correct assessment of the surface texture is often hindered by the velipellis which can result in an unusual development of the underlying pileipellis. also aberrant surface due Old specimens may show an pileus to disruption on ageing, causing a secondarily squamulose to squarrose aspect. Microscopical ex- amination of the pellis, however, will usually reveal its true nature. 3. Lamellae The lamelllae are broadly to narrowly adnate, sometimes even (almost)Tree. This character is hardly applicable as an infrageneric character, although species considered as adnate lamellae. relatively primitive usually possess broadly Young lamellae are often but violaceous white, yellow, grey or tinges are sometimes present. Lamellae soon turn brown as the spores mature, but sometimes this process is rather slow. 2 K u Y p E r: Inocybe in Europe I 3 Much value has been attached to the presence of olivaceous tinges in the lamellae, but in my opinion incorrectly so. The colour of the lamellae in mature specimens is determined five different viz. of intracellular in by factors, (i) presence pigment the colour of of basidia lamellale, (ii) ripe spores, (iii) presence necropigmented of (in subg.