The tribe Leucopaxilleae was recognized by Singer (1948: 29) to be a group of related genera within the Tricholomataceae, characterized by the tricholomatoid or clitocyboid basidiocarps, non-hymeniform epicuticular layer of the pileal surface, and the production of amyloid basidiospores. Prior to the use of the amyloid reaction of the spore as a primary character for classification of the Agaricales, the species of the Leucopaxilleae were to be found amongst such diverse genera as Clitocybe (Fr.) Kummer, Lepista (Fr.) W. G. Smith, Omphalina Quilet and Tricholoma (Fr.) Kummer of the tribe Clitocybeae (Tricholomataceae), Collybia (Fr.) Kummer of the tribe Collybieae (Tricholomataceae), Hygrophoropsis (Schroet.) Maire and Paxillus Fr. (Paxillaceae), and Cantharellus Adanson ex Fr. (Cantharellaceae). At the present time the Leucopaxilleae contains eight genera recognized by Singer (1962: 142), of which all, except the monotypic North American genus Pseudoarmillariella (Singer) Singer, are known from the British Isles. Lentinellus Karsten was initially included on the basis of the amyloid verrucose spores but this genus is now generally accepted (see Maas Geesteranus, 1963: 430; Donk, 1964: 246) as being more closely related to members of the Auriscalpiaceae in the Aphyllophorales, showing similarities in the dimitic hyphal system and the presence of gloeocystidia. All species in the Leucopaxilleae are characterized by amyloid reaction of the spore wall to Melzer's reagent. It is a constant reaction, although initial treatment with ammonium hydroxide may occasionally be necessary in older collections. All species of Melanoleuca Pat. and those species of Leucopaxillus Boursier with ornamented spores show a very strong, blue-black reaction of the verrucose ornament to Melzer's reagent, whilst the smooth-spored species of Leucopaxillus, e.g. L. giganteus (Sow. ex Fr.), Singer give only a weakly amyloid reaction. The amyloid reaction is due to the presence of either an entire or a fragmenting perisporial layer. Josserand (I940: 21) demonstrated for both Leucopaxillus and Melanoleuca the existence of an ornament ('ornementation fondamentale') below the fragmenting perisporium, by removing the amyloid material with sodium hydroxide solution. However, although the underlying ornament may be well developed in spores of the Russulaceae, in Melanoleuca it was described as insignificant or non-existent. Nevertheless, such a structure was clearly demonstrated by Perreau-Bertrand (I967a: t. II H) for M. melaleuca (Pers. ex Fr.) Murr. (as M. vulgaris Pat.) using ultrathin spore sections in which a full complement of wall layer was visible, even though the wall appears very thin when observed with the light microscope.
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