FaARLOW RererRence USRARY ISSN 0374-7859 ss NaS oo sep 0 4 1982 ) THE GARDENS’ BULLETIN SHEE aS SINGAPORE ( ae GARDENS’ BULLETIN SUPPLEMENT NO. 2 1991 i EN a Xe x , Trogia (Basidiomycetes) J By - % EJ.H. Corner, CBE. ERS. \ Emeritus Professor of Tropical Botany University of Cambridge » § \ a () () Published by \y 2 National Parks Board ( Singapore Botanic Gardens N4 Cluny Road 0 Singapore 1025 N ‘ aj y OO RRODER EOE Trogia Trogia (Basidiomycetes) BY EH. Comet. CBE. ER:S. Emeritus Professor of Tropical Botany University of Cambridge Published by National Parks Board Singapore Botanic Gardens Singapore 199} Sold by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 1025 For sale only: not for exchange Price S$17 © National Parks Board 1991 Printed in Singapore by Letraprint CONTENTS PAGE (PURGE KCRG San candecncecbeccdaceunee eeaneedn:2 cua AO Se REESE APICES een Ae re) ie ] MEDEA ONS By ESTOS weal oe ae ae eS OC a 2 PS HO ACC AC ee VR Ne Te IM OO er ofa sod Sock eed ave Neacsadudeasddeccatmcenchaietaasse: 4 GRY SHIPS spancsnecccothsscecteete ite aC ORp eee SSS EEO ET EEE Cn EE Pe er SCR oe ee 5 NACI EATOIA GCS mene OEE eee POO RN nM: oD. 2 sion Gace oGs nena samnleebeenesas sewesdsbues oes 6 'RENERENCSS. s-cuoncsacondanebecenadoe bene Rene eee ee ene a ie Rn eo 6 Mera ineirapical Asia: and Australasia ..i3s2.sc2ic.de55. Seskeece ds veh ewcssadeevasbessveesseees a) Key to identification on macroscopic characters ..............00ccccccccceceeseeeeees 7 ey tO identiication ON MICTOSCOPIC ChALAaCtEMS’ ...............2000...sseeseeceeenede oes 10 SPECIE GeserIplions (alphabetiCal Oden) as._.2.. .fenris.+-0s<00..susasonaocecececeeness 16 2 OE Re TGC UIN STENCH oe oc yea ne 87 Nr ACUMEN HRT EATON feo Pes PEt YS oS oie am a! 87 Beceiierdescriplionss (alphabetical OLGEL)” .csc.ccfccc2e.8. ccs. v ened oabeeccetecseseen socks 88 EAU DOLD TAA LTUG! ens ss Reece RESIS AEE CRE a 93 OSISCHIPIR (AGRE LELEMCES) I cass se eee tence ae nk ss cts decccnne cosceeseeceveeesersecss 97 Digitized by the Internet Archive in 2014 http://archive.org/details/gardensbulletins42unse Trogia (Basidiomycetes) E.J.H. CORNER 91 Hinton Way, Great Shelford Cambridge CB2 SAH England EFFECTIVE PUBLICATION DATE: 23 MAR 1992 Abstract 73 species (38 new) of Trogia are recorded from tropical Asia and Australasia, and 17 (3 new) from the neotropics. Keys to identification are given and, for most species, descriptions. The sarcodimitic fruit- body is regarded as a refinement to the lignicolous habitat, and tending, consequently, to simplification by loss of stem and gills, as well as by diminished size. Its origin is considered along with the polypore Meripilus and a possible explanation is found in the ancestry of pteruloid fungi with inflating hyphae. Heimiomyces, with marasmioid affinity, is compared. Rimbachia is considered along with Rhodoarrhenia (one new species) and Skepperiella. New species — 7rogia aurantiphylla, T. basivillosa (neotropical), T. brevipes, T: carminea, T. decipiens with var. pleurotella, T. delicata, T. diminutiva, T. endocystidiata, T. exigua, T. fusciceps, T. fuscoalba with var. metuloidea, T. fuscolutea with var. minor, T. icterinoides, T. laeta (neotropical), T. lateralis, T. latifolia, T. limonospora, T. limonosporoides, T. macra, T. mamillata, T. marasmioides, T. minima, T. nigrescens with var. violascens, T. nitrosa, T. obfuscata, T. octava, T. odorata, T: omphalinoides with var. confertifolia, T. pallida, T: perpusilla (neotropical), T: polyadelpha, T. pusilla with var. sublateralis, T. raphanolens, T. revoluta, T. rosea, T. seriflua, T. sublateralis, T. subtomentosa, T. subtranslucens, T. tenax, T. venulosa — Rhodoarrhenia solomonensis. Other new varities — Trogia alba Corner var. brasiliensis and var. minor, T. anthidepas (Berk. et Br.) Corner var. brasiliensis, T; aphylla Corner var. solomonensis, T. fulvochracea Corner var. brasiliensis and var. solomonensis, T. impartita Corner var. griseola and var. major, T. subglobospora Corner var. mellea. New combinations — Trogia guadelupensis (Heim) with var. nov. subincarnata. Introduction My treatment of this genus in 1966 brought an outburst of indignation. That anyone should assemble in one genus agarics with and without gills along with stereoid or sessile forms on the grounds of a sarcodimitic construction was considered a farce. I replied (Corner 1968) and the matter has rested until the recent contribution by Redhead (1987). I am not deterred because I have had numerous other collections of these fungi, on which I now report. They link even more closely the variety that is to be found in this alliance and confirm my understanding of the genus. According to Singer (1975), the species which I put in 7rogia should be dispersed in Gerronema, Hemimycena, Hydropus and Mycena, to which one can add Marasmiellus and Neoclitocybe. His reason is that what he refers to as the supposed sarcodimitic construction occurs widely in Tricholomataceae and, if accepted as a generic distinc- tion, then 7rogia would be augmented by many more species. I agree and look forward to the revelation because I have no faith in those genera, unless in Mycena. I cannot discover any clear distinction between them, even though dispersed in three or four tribes, and I doubt their natural content when I find such as the simply monomitic Omphalina fibula and Cantharellus albidus placed in Gerronema along with species that have the sarcodimitic structure of 7rogia; when the common Trogia infundibuliformis, of wide acceptance, is described as a new species of Neoclitocybe, namely N. mem- branacea Sing. et Grisl. (Pegler 1977); when 7: cantharelloides is still maintained in 7 TROGIA AS A GENUS Trogia, though it is, as originally described, a species of Panus (Corner 1981); and when Vanromburghia is unexplained. Indeed the concatenation of species in Trogia is far closer than that which leads to the inclusion of Cyphellostereum in the same tribe as Tricholoma. In practice, as I have ascertained by studying hundreds of different agarics, the sarcodimitic construction is easily recognised and extracts a close alliance of agarics with a characteristic waxy-cartilaginous appearance that is readily detected in the field. If a strip of stem-tissue is detached from a living specimen, or from one preserved in alcohol-formalin, and teased apart, the very long and more or less thick-walled fusiform cells can be seen at once under a moderate magnification. It is a very useful guide to identification because it extracts from the welter of collybioid, clitocyboid, mycenoid, omphalinoid, pleurotoid and tricholomatoid agarics and gives precision. Indeed, sooner or later in the course of specific identification, it is necessary to examine the hyphae and, as their behaviour characterises the fruit-body, so I regard it as a primary con- sideration. My first enquiry on studying any fruit-body microscopically is to find out what kind of hyphae make it. I note that with dried material only of Trogia, this may be impossible because the hyphae often stick firmly together; hence, doubtless, the neglect of the sarcodimitic construction. It required the passage of thirty years before the importance of hyphal construction was realised in connection with polypores. The species of 7rogia are abundant lignicolous fungi in South East Asia, probably throughout the tropics, and they come at once to the notice of the mycologist and ecologist concerned with decomposition on the forest floor. Though the fruit-bodies are usually small, they are often densely gregarious or caespitose and may occur in large troops on fallen trunks. They prefer, it seems, wood in the later stages of decay and, as it may then moulder into soil, so some species appear to be terricolous, e.g. T. cyanea, T. delicata, T limonosporoides, T: pallida and T: subgelatinosa. The largest fruit-bodies that I have met are those of 7. fuliginea and T. /atifolia with the pileus up to 11 cm wide. The smallest are those of 7) minima and T. perpusilla with the pileus 1-3 mm wide. I list about 73 species for tropical Asia and Australasia and an under- estimated 17 for the neotropics. There are certainly more to be found and many im- provements to be made, but the keys and descriptions are the initial guides. In the monograph (Corner 1966) only Latin descriptions were given for new species. These I have translated in the following account but, where the description was in English, I have referred to that monograph (with figures and plates), unless later collections have enabled me to improve. The identification of the species with white or yellowish fruit-bodies is intricate because they are variously depigmented and reduced from several alliances; recourse to microscopic detail is inevitable. Rhodoarrhenia is an uncertain genus which holds some species allied with Trogia. Trogia as a genus In spite of the criticism that I have lumped too many different kinds of fruit-body into this genus, I am still unable to find any distinction sharp enough to separate them into genera; there are in all cases intermediates and they cannot be ignored. Modern taxonomy imposes humanistic limits which are shown, sooner or later as mycological discovery progresses, to be unreal; its track is strewn with waste genera. In 7rogia the one isolated feature is the asperulate spore of 7: pleurotoides. The asperulate spore, as opposed to the smooth spore, is often taken to distinguish two genera without con- sideration why the spore should be asperulate. In the case of Boletus and polypores, I have found evidence that, as one might expect from its predominance in modern basidiomycetes, the smooth spore has evolved from the ornamented and bitunicate (Corner 1972, 1989). Ornamented and smooth, or nearly smooth, spores are admitted for instance, in Jnocybe, Lepista, Melanoleuca, Ramaria and some genera of polypores. Therefore, the asperulate spore may be expected in 7rogia. INDECISIVE CHARACTERS 3 The ideas of Vanromburghia, based on Trogia silvestris, and of Inflatostereum, based on a stereoid species of Trogia, are also vapid. The first is a mesopodal Trogia without gills, but there may be traces of gill-folds about the stem-apex, and such gill-folds are better developed, though also disappearing, in the allied 7? rosea, T. pallida and T. macra (as here desribed), and similarly in 7 aphylla. There are enough microscopic details among these five species to split off three microgenera, if no others were known, but all link up with fully lamellate alliances. Gill-less species occur, for instance, in Laccaria and the alliance of Marasmius without generic effect. The fruit-body of Inflatostereum is not only devoid of gills but is pleuropodal or sessile and unilaterally flabelliform; microscopically, it is toughly sarcotrimitic with thickening hymenium; thus, it appears stereoid except for the inflating hyphae and, in T. rivulosa, the slightly incurved margin of the pileus which is a relict agaric feature. Several species of Trogia, as given in the keys, have variously simplified and narrowed gills to excentric, sublateral and lateral stems, and among these 7. pleurotoides approaches in form and colour what one would expect as ancestral to 7. stereoides. In parallel, there is the transition from mesopodal 7. infundibuliformis through T. venulosa and T. subtranslucens to the stereoid 7. cervina. The idea of Inflatostereum seems to have been achieved twice by Trogia. The sarcotrimitic construction is incipient in the lamellate 7 obfuscata. The thickening hymenium occurs in the gill-less 7. silvestris, the partly gilled JT rosea and T, pallida, and in the lamellate 7. /ateralis and T. pleurotoides where it is most con- spicuous in the gill-intervals. Thus, with loss of gills, the thickening hymenium may compensate loss in spore-output, if that is material. The thickening hymenium appears to be a primitive feature in basidiomycetes and traces may be expected in Trogia. Indeed, I see in Trogia a genus no more varied than Thelephora with its transition from fully clavarioid forms to mesopodal pileate, stereoid and fully resupinate, with or without the pseudo-hydnoid hymenial papillae; the reason is that there are many species bridging these humanistic distinctions. A natural genus is found, not dictated. As some indecisive characters in 7rogia, I note the following: — 1. Pileus-shape, whether plane, umbonate or umbilicate to infundibuliform. These differences can be specific but several species are variously intermediate. Why the pileus should be umbonate in 7rogia is not known. In terricolous agarics it com- monly results from the subterranean origin of the fruit-body with conical pileus thrusting through the soil, but this cannot be the case with lignicolous fruit-bodies of superficial origin. 2. Lateral stem. This distinguishes some species but in others the stem is sublateral in that there is a small development of the pileus on the side towards the sub- stratum. Some species are variable in the position of the stem, e.g. 7. decipiens and 7. pleurotoides. 3. Gills, whether well-formed or pliciform. The first state seems to be the distinction between Gerronema-Hydropus and Trogia as originally defined but some species have both kinds of gill in the same pileus, e.g. 7) inaequalis (Pegler 1986), 7. macra, T, pallida and T. rosea. In the fully laminate species the gills may be narrow in some to fairly wide or very wide in others. Some care is needed, however, in the use of this feature because the gills may begin to function when narrow and, then, widen considerably by intercalary growth of the hymenium. In so doing, the at- tachment of the gill may change from decurrent or adnate to sinuate and ventricose, even adnexed, e.g. 7. fuliginea, T: revoluta. This widening, of course, thrusts the limb of the pileus upwards and alters its shape. 4. Spores, amyloid or not. The reaction may be so weak that it is impossible to decide, especially if some glycogen inside the spore gives a slight vinaceous brownish colour, e.g. 7) aphylla var. solomonensis. In T. straminea the spores may or may not be slightly amyloid. 7; impartita has pale amyloid spores but they are inamyloid TROGIA AS A GENUS in the very similar 7, raphanolens; likewise with the species-pairs 7. omphalinoides- T. octava and T. tricholomatoides-T. latifolia. In T. alba the spores are not amyloid but slightly so in its var. minor; similary, 7) ceraceomollis and its var. amylospora. The amyloid reaction seems to be caused by a very thin coat of mucilage as a remnant of an exospore. . Cheilocystidia, as a sterile gill-edge or not. The variability of this feature is shown by 7. subglobospora with sterile, partly and wholly fertile gill-edge. In some species the cheilocystidia collapse early, even before sporing has begun, and mature fruit- bodies appear to lack them. . Pleurocystidia, present or absent. Their presence is a useful mark for numerous species but in others they may be so sparse as easily to be overlooked, e.g. 7 calyculus, T: polyadelpha and T. subdistans; they may even be as evanescent as the cheilocystidia. Then, 7: /imonosporoides has pleurocystidia and its ally 7. /imono- spora lacks them; similarly 7) fuscoalba with T. subdistans and T. seriflua with T. icterinoides. The thick-walled pleurocystidia of 7: fuscoalba var. metuloidea suggest a good distinction but they are slightly thick-walled in several species, e.g. T fuscolutea. . Pileus-structure, with or without a pseudoparenchymatous hypodermis. This is also a useful character but it may be confined to the centre of the pileus and it is variable in the complex of 7? umbrino-alba. Generally, the rugulose surface of the pileus results from this hypodermis. . Pileus-surface, with or without ‘rameales structure’. Such a surface is well-developed in 7: marasmioides and T: subtomentosa, \ess so in T. venulosa and vestigial in its ally 7. infundibuliformis. . Fusiform cells. These are the long, inflating and, usually, thick-walled cells which, in my monograph, I called the skeletal cells; they certainly support the fruit- body. They may be limited to the stem or developed throughout the fruit-body or variously extended into pileus and gills. They vary in length and width in every species but in some they are noticeably very wide or long. It would be desirable to test specific alliances by the maximum length of the fusiform cells but this is difficult to ascertain because the longer are nearly impossible to extract without breaking. I have not used the feature in the analytic keys. . Sarcotrimitic construction. While this distinguishes most stereoid species, it is present in varying degree in the stem if not also in the pileus, of some lamellate species, e.g. 7. obfuscata. Specific alliances. These are given under the specific descriptions. I note the following more extensive alliances but, in the generality of the construction of the fruit-body and in the uncertainty of the limits of the alliances, they are not separable as genera: — Ne 2 3: 4. With bluish green or fuliginous green fruit-bodies — 7. odorata, T: subviridis, T. cyanea, T. pleurotoides and T. stereoides. With lilaceous fruit-bodies without cystidia — 7. infundibulifomis, T: venulosa, T. subtranslucens and stereoid 7. cervina. With pleurocystidia — 7. aurantiphylla, T. fulvochracea, T: fuscolutea, T. silvestris, T. rosea and T. macra, both the last with narrow gills or none. Without pleurocystidia — 7: furcata and T. pallida. Mycenoid — T. delicata, T- omphalinoides, T. revoluta. With endocystidia — 7. endocystidiata, T. limonospora, T. limonosporoides. Xerulaceae In his recent contribution to the problem of 7rogia, Redhead places the genus in a primitive position in the family Xerulaceae which has, as advanced members, MERIPILUS 5 Oudemansiella and Mycena. His thesis is that the sarcodimitic construction advances, or simplifies, into the monomitic with inflating hyphae as in Oudemansiella and the inflating but secondarily septate as in Mycena. Inflatostereum may be a primitive stereoid member of the family. Thus, 7rogia is supposed to be in the process of developing stem, gills and centric pileus. This is quite contrary to my original view that Trogia displayed the degeneration of the agaric into the gill-less and stereoid state (Corner 1966). I do not find this Xerulaceous argument at all convincing; a morphological series can be read in either direction and time’s arrow is needed for evolutionary sequence. The essence of the sarcodimitic state lies in the specialisation of tissue enlargement by means of the fusiform cells and not in the conjunction of ordinarily enlarged and unenlarged hyphae, for that is a monomitic construction, as I illustrated for Clavulinopsis (Corner 1950). The clavarioid comparison is seen in Prerula which, if its long skeletal cells inflated, would be sarcodimitic. I note that Redhead’s inclusion of Lentinus tigrinus in the sarcodimitic category is misleading (Corner 1981). In complaining as others have done, that I have lumped too many genera in 7rogia, Redhead prunes off without explanation how this can be done and overlooks the bridging species. What he has left in 7rogia is not clear. In any case, the fruit-bodies of this genus with thin flesh, hollow stem, and small spores and basidia cannot be antecedent to those of Oudemansiella with thick flesh, massive and solid stem, large spores and large basidia. The. construc- tion of 7rogia reveals specialisation and not primitive evolution among its modern species. Nevertheless, in his valuable contribution, Redhead indicates sarcodimitic species in Baeospora, Clitocybula, Mycena s. |., and Xeromphalina. From my studies of English species, I add Mycena crispata Kuehner and M. delectabilis (Peck) Kuehner, both of which are placed in Hemimycena by Singer. They are small white species which would come in Group D of my key on microscopic characters and near to 7. octava and 7. exigua respectively. The interest lies in discovering how the temperate species of Trogia fit in with the tropical. Meripilus This genus of polypores introduces a problem which, if it can be solved, will lead to much better understanding of basidiomycetes. The fruit-bodies have the sarcodimitic construction but, so far as known, consistently without clamp-connections (Corner 1984). Either Meripilus is a parallel evolution from a monomitic source of polypores, such as the ancestry of Grifola might supply, or it is connected with the ancestry of Trogia. My preference is to think of the pre-Preru/a and pre-Deflexu/a with inflating hyphae whence an extinct hericioid ancestor might have diverged into the poroid Meripilus and the lamellate Trogia. The only evidence that I can find for such a hericioid ancestor lies in the microfibrillar flesh of M. giganteus, as if the fruit-body were com- pounded of clavarioid branches. Parallel instances are Panus s. str. and Rigidoporus with uninflated hyphae and Lentinus s. str. and Polyporus s. str. with their peculiar hyphae. Thus, I differ profoundly from Redhead’s view that /nflatostereum evolved into Meripilus and Trogia. He gives no explanation how tubes (or gills) were evolved and disregards the difference in basidia and spores in the case of Meripilus. That tubes may variously degenerate in resupinate fungi with uninflated hyphae is no proof that they were originally so easily evolved. Moreover, the idea that a stem is an advantage in lignicolous basidiomycetes is not borne out by Hymenochaete, Stereum s. |., Thelephora and polypores in general, or by the great abundance of resupinates. Already raised on woody supports, the fruit-body does not require a stem. It is to be noted that, in many species of Trogia, the caulocystidia and pileocystidia form a close palisade or hymenioderm in the unexpanded fruit-body. Then, on expan- sion, the layer becomes disrupted into the pruina-like clusters on the stem, the 6 HEIMIOMYCES furfuraceous centre of the pileus, and the scattered pileocystidia over the limb. That such a hymenioderm is the result of reduction from an original trichoderm is shown by diverse genera, e.g. Boletus, Entoloma, Pluteus and Amauroderma. It is part of the sterile cortication of the ancestral fruit-body, and the loss of the trichoderm is shown in Meripilus. Thus, Trogia marasmioides and T. subtomentosa are relatively primitive in this respect, parallel with 7richolomopsis. The state with smooth pileus, lacking cystidia, is the advanced. So Oudemansiella and Xerula are more primitive in this respect than most species of 7rogia. Heimiomyces The type-species of this genus, H. tenuipes (Schw.) A.H. Smith, is a frequent pan- tropical fungus with which I have been acquainted since 1929, but it was not till some years after I published the account of Trogia that I realised the Trogia-like construction of the stem. The inner tissue of the hollow stem has long, fusiform clamped cells 8-28 pm wide, with thin or slightly thickened walls. They become subdivided by many broad clampless secondary septa into cells 60-750 ym long. Unfortunately, I have not had material young enough to reveal at what stage secondary septation begins or how long the initial fusiform cells may be. Among these inflated cells there are many uninflated hyphae, 2-5 um wide, with thickening walls and often branched from the clamp; thus the stem is approaching the sarcotrimitic state. In contrast, the hyphae of the pileus and gills are monomitic and uninflated with more or less mucilaginous walls. Redhead gives the species, under the name Xeromphalina tenuipes, as sarcodimitic ‘in stipe, modified’. Thus, Heimiomyces might be regarded as a trichodermatous Trogia. However, H. tenuipes has the acerose basidioles of Marasmius, which are absent from Trogia, and the peculiar uninflated gelatinous tissue of the pileus which is also found in the alliance of Marasmius. Until such fungi have been analysed more fully, I prefer to keep Heimiomyces distinct. Xeromphalina lacks the acerose basidioles and has monomitic short-celled hyphae with broad septa, inflating in all parts of the fruit-body, as in Flammulina and Omphalina, though Flammulina may have some secondary septation in the stem. References Corner, E.J.H. (1950). A monograph of Clavaria and allied genera. Ann. Bot. (London) Mem. 1. . (1966). A monograph of cantharelloid fungi. Ann. Bot. (London) Mem. 2. . (1968). Mycology in the tropics. New Phytol. 67: 219-228. . (1972). Studies in the basidium. Gdns’ Bull. Singapore 26: 159-194. . (1981). The agaric genera Lentinus, Panus and Pleurotus. Nova Hedwigia, beth. 69. . (1984). Ad Polyporaceas III. Nova Hedwigia, beih. 78. Pegler, D.N. (1977). A preliminary Agaric Flora of East Africa. Kew Bull. Additional Series VI. . (1983). Agaric Flora of the Lesser Antilles. Kew Bull. Additional Series LX. . (1986). Agaric Flora of Sri Lanka. Kew Bull. Additional Series XII. Redhead, S.A. (1987). The Xerulaceae (Basidiomycetes), a family with sarcodimitic tissue. Can. J. Bot. 65: 1551-1562. Singer, R. (1975). The Agaricales in modern taxonomy. J. Cramer; Vaduz. Ie Trogia in tropical Asia and Australasia KEY TO THE SPECIES ON MACROSCOPIC FEATURES . Gills absent or as slight folds disappearing in the outer half of the pileus ............. Group A . Gills present, not so disappearing. DRE TCULO POM hae eee et ee oe ee ee ee ee tls Se ES 2 Group B p. 7 2. Mesopodal, varying excentric in some species. 3. Pileus orange or red, mostly small, —10 mm wide. 4. Pileus and stem orange. Gills decurrent, ochraceous. Ceylon ........ T. inaequalis p. 4. Pileus rose red. Stem white. Gills adnexed, pink. Borneo ............ T. carminea p. 3. Larger, differently coloured. 5. Pileus pinkish vinaceous, then pallid. Gills narrow. 6. Stem becoming hollow with the pileus pervious .......... T. infundibuliformis p. Gee SLEMMNOL NOMOW PM SOLOMONMIS 6: ccecc os cs oie; oxo cy 2a res 52 aye ora be a ose weaniogs T. venulosa p. 5. Fuliginous green to indigo (see below). 7. Gills —0.5 mm wide. Mesopodal to pleuropodal .............. T. pleurotoides p. 7. Gills 1-3 mm wide, subdistant. Se Smells Olranisceds SOLOMON NSlevas lc sce vs cee ee secs ss ee T. odorata p. 8. No such smell. Flesh of pileus agglutinated. Malay Peninsula wu ate Hac Ny ere ee eb ORR RR OES acl eee T. cyanea p. 4] 21 41 85 64 60 24 5. Differently coloured. 9. Stem or gills yellow. Pileus yellowish, often clouded fuscous or fuliginous, mostly concave tonmiundibulitorme: Gills mostly dectitrent’ [22> 2.5. 4--¢- 2 s5s464- oe Group C p. 8 9. Gills and stem not yellow. LOSeieuss brown tO 1nueiMOus- OF Prey ee cme ese reece aes eee see Group D p. 8 LO Pilevuscwinteron yeuowish) winter eee nie. see cc esses ec es Group E p. 9 Group A 1. Pleuropodal, flabelliform ascending, shortly stipitate to subsessile, stereoid. 2. Fuliginous olivaceous, at least towards the pileus margin ................. T. stereoides p. 73 DRmuUSCOUS WIth PICY NYINEMIUM cris noon oe acces ocee eee ee sete eee eres T. rivulosa p. 69 2. Brownish to ochraceous tan, sometimes with a mycelial skin spreading over the wood eee a a re a= ae ws (ade cree sais eat state a tale cee lece Serie ood T. cervina p. 23 PerGrevish tinsed: vinaceous, subiransiucent ..2-. 0... eee acstaa esse eee T. subtranslucens p. 79 DMVIMILC EVEN ESIIANL | GENCALG panacea nte Seas soa ta aaa ah oon Dias eee T. minima p. 52 Mesopodal or, if pleuropodal, with slender stem (7: aphylla). 3. Pileus usually papillate-umbonate and rugulose. Ara ENEUSEGUIMSON tOLrOSE-PID Kare seen an oe mee Se oem e ete a eee eeeeese T. rosea p. 69 4s Pyleus cinnamon: fawn to pale @chraceous =---=---.--st-- sesh reese mee T. silvestris p. 73 3. Pileus neither papillate nor rugulose. 5. Fuscous olivaceous to fuliginous yellowish, mostly without gills. Stem 0.5-1 mm thick F OCIS BACON a ea Paria Enc O ais AC Stock eieain isla aetna ie ea ae Berar T. aphylla p. 18 5. Pileus pink, yellow or white, mostly with slight gill-folds. 6. Pileus —9 cm wide, pale clear yellow. Stem 2-4 mm thick .............. T. macra p. 47 Semstera d—lsSimpmisthicks cece eek ores coke cee Se hue GER ee T. pallida p. Se. Stem O05 smmbthicks Pileus+2—=ssmm wide. .secencereee es - T. diminutiva p. Group B IeeBuliginousspreenito indigo Gillsiveny narrows. 4: sce" cnrresemiels seems ce T. pleurotoides p. i= Pileusefascous: brownish. iGills: ochraceous... + pee seme 2s see es cree T. ochrophylla p. 6. Pileus smaller, not yellow. 7. Pale livid pink, subochraceous with age. Stem —8 x 2-3 mm, short Saree oeare Sore S SSG Catan nS eke Ne ae Sis SMO ee Pte: areeyen: ihre’ ser. T. venulosa p. 7. Pallid white. 8 TROGIA IN TROPICAL ASIA AND AUSTRALASIA 1. Yellowish white to white. 2. Stem sublateral, brownish at the base. Gills 0.5 mm wide ............... T. sublateralis p. 77 2. Stem truly lateral, not brown at the base. Se Pileus#1=3imiiwides:Gillstasi2—4 afaintefoldss eee eerie eee T. minima p. 52 3. Larger. 4 Gills: —2.mmi Wide... ..c.. cee es Hod at eee eee T. lateralis p. 41 4 1Gills#2—44mmy wide: Stem vanyingacentrall ss. see einer eine ites T: marasmioides p. 50 Group C 1. Gills adnate to adnexed, mostly with clear yellow edge. Pileus papillate-umbonate. 2 GillssoraneestOsnichnyellowsSOlOn One| S lamer aereeteene er rae ee T. aurantiphylla p. 18 2. Gills white. 3. Gills —4 mm wide, edge yellow. Pileus with fuscous centre ............ T. fuscolutea p. 36 3. Gills —1.5 mm wide, not yellow. Pileus and stem yellow ................ T. papillata p. 63 1. Gills decurrent. Pileus not papillate-umbonate. 4. Small. Pileus —20 mm wide. Stem —1.5 mm thick. Gills —2 mm wide. 5. Pileus not fuscous, —12 mm wide. 6. Pileus primrose yellow, mesopodal. Gill-interstices subrugulose ...... T. primulina p. 65 6. Pileus pale ochraceous, excentric. Gills not veined. Stem brownish at the base Seep A IG Sr AIG oes OOS SIS Sel oiata Soste ho ake a Grater T. sublateralis p. 77 5. Pileus fuscous in the centre. Gill-interstices not or scarcely veined. 7. Pileus —7 mm wide. Gills 0.5 mm wide, whitish. 8. Basidia 2-spored. Spores 7-9 x 5.5-7 um. Stem central to sublateral 3b gubuish guev'ee pod Sy feSeee iw Lglstenay tee aT isn ey egret del aa ow ATA e ee eR Oa Meee ee T. pusilla p. 66 82 ‘Basidia4-sporedsspOresy4—siy puri Oy Dayana nee een eee T. fuscomellea p. 37 7. Larger. Gills —2 mm wide, honey-yellow. Basidia 4-spored. Spores 7-9 um long ee ere ONIN CaS OS e Ie AeA oo Sa gn os Mae oe oe a lr. mellea p. 50 4. Larger in all parts. 9. Pileus and stem greenish yellow. All parts reddening with potash. Ceylon eee Om Ie eras COR SOG ONGOes Gt.c OOOOH IDC ROC OU SOO ne T. holochlora p. 38 9. Not so. Gill-interstices often veined to poroid-reticulate. Pileus fuscous to yellowish. Stem and gills yellowish. 105 ‘Spores; subeloboses4—s- 5a eer re ere tee eee T. subglobospora vy. mellea p. 77 10. Spores ellipsoid. 11. Pileus with dark innate fibrils. Stem fuliginous pruinose ...... T. anthidepas p. 17 Is Without such fibrils on proinalesere eerie cee a eieeernein T. straminea p. 74 (Rileus with) tomate fib mils ieee eee ere ee eee eee T. aff. straminea p. 75) (Stem white, yellowish in age. Gills grey then white ........ T. umbrino-alba p. 83) Group D 1. Gills adnexed, sinuate or adnate, rather wide, rather crowded. 2. Smell@strones mitrouss Etleus iDLOWAlISHi ee areata eee niente er eee T. nitrosa p. 56 2. Not nitrous. 3. Gills 10-20 mm wide. Pileus 5-11 cm wide. 4. Pileus acutely umbonate, yellowish white clouded fuscous in the centre. Ceylon eT eer ee ITS ao 25 6.6 OGRE tks op Gut T. tricholomatoides p. 83 4. Pileus subumbonate, pale fuscous brownish. Borneo ................. T. latifolia p. 43 3. Gills not so broad. Pileus not so large. 5. Pileus finely fibrilloso-squamulose to subtomentose, pale brownish subochraceous with fUSCOUS A COMUNE lore iss 5 te 2e10.0:4. 60 epereire fe Lee epee Ee TERE T. subtomentosa p. 79 5. Pileus not subtomentose. 6. Pileus acutely umbonate, cinnamon ochraceous to fulvous fawn or fuscous grey. Stem 11) on ee oO eee Ce Orn St a ten Oe eA oo OS T. fulvochracea p. 31 6. Pileus umbonate or not, of dull colour. 7. Pileus greyish lilaceous. Stem 3-7 cm, —9 mm wide at the base. Smell farinaceous, strong. Gills) =—7omm wide’.s..-- +o. oe aoe eee ae ee T. lilaceogrisea p. 44 KEY ON MACROSCOPIC FEATURES 3) 7. Pileus fuscous umber to greyish. Stem smaller. Gills usually narrower. 8. Pileus, stem and gills hispidulous. Smell farinaceous or none SBC Mee eee ee ae ee ne ee eee T. hispidula p. 38 8. Pileus not hispidulous. 9. Pileus rugulose. Stem white. Gills greyish Shs HES RE CaS ao er eee T. fulvochracea vy. fuscogrisea p. 32 9. Pileus not rugulose. Stem fuliginous pruinose. Gills white JaSR ART Re ease Ree 22 eS roe ne T. mycenoides p. 53 1. Gills more or less deeply decurrent. 10. Stem sublateral, short. Pileus —25 mm wide, cinnamon brown to somewhat orange or ferruginous + Se ResSobgeme eS eh Ae Oh eee A eet ee eee ee T. tenax p. 81 10. Mesopodal. Pileus more or less infundibuliform. 11. Gills narrow 0.5-1 mm wide. Fruit-bodies small. 12. Pileus —6 mm wide, honey-yellow with fuscous centre. Stem yellow ... 7: fuscomellea p. 37 12. Pileus and stem not yellow. 13. Gills forked 1-3 times, 0.5 mm wide, subdistant ............... T. furcata p. 33 13. Gills not or rarely forked. 14. Pileus —11 mm wide, grey with innate grey-green streaks .. 7. calyculus p. 21 12> Pileystlarper: LuSCOUS UMIDEn..-c - .- - Soee es earn as os T. umbrino-alba p. 83 11. Gills wider. 15. Pileus with greenish or bluish tints. 16. Pileus fuliginous cyaneous. Gills and stem paler ............... T. cyanea p. 24 16. Pileus grey tinged greenish ochre. Gills and stem white ....... T. subviridis p. 80 15. Pileus not so tinted. 17. Stem fuscous or fuliginous pruinose. 18. Gill-edge fuscous fuliginous pruinose as the pileus and short stem a ee te SE ae ee INE corer e tO DE, cla ci anesa T. brevipes p. 20 18. Gill-edge not darker pruinose. 19. Gills whitish, —2.5 mm wide. Stem whitish .......... T. fusciceps p. 33 19. Gills —6 mm wide, paler concolorous with the pileus SE Se ee tae eas eles T. ceraceomollis p. 22 17. Stem whitish pruinose. 20. Uniformly grey, firmly waxy subgelatinous. Pileus —7 cm wide. Gills 2.5-6 mm Wider Ree Saree: ce. RES ES ee wat T. subgelatinosa p. 75 20. Not so. 21. Spores subglobose 4-5.5 x 3-5 um. Gills 1-2 mm wide, white, interstices OVO) Gaeta ee a rays J Soa so T. subglobospora p. 76 21. Spores ellipsoid. 22. Spores 4.5-5.5 x 2.7-3.3 ym. Gills —2.5 mm wide, thick, distant, white ERICH LUSCOUS Wo seer ee a= Baca e ieee ciara es weiner = T. obfuscata p. 58 22. Spores larger. Gills 2-4 mm wide, interstices not or scarcely veined. 23. Pleurocystidia thick-walled .... 7: fuscoalba v. metuloidea p. 35 23. Pleurocystidia thin-walled. 24. Spores 8-10 x 6-7 um. Stem greyish .... 7. cystidiata p. 25 24. Spores 6-8 x 4.5-5S.5 um. 25. Gills 2-4 mm wide, subdistant ..... T. subdistans p. 75 25. Gills —2 mm wide, crowded, hispidulous. Pileus papillate- UNI bONALC eet sae5 sere SS evens oe T. fuscoalba p. 34 Group E 1. All parts of the fruit-body blackening. Flesh with watery juice turning red or violaceous on exposure. Gillsvadnate;1—2 mm: wide; very crowded. 2.7252 ce te 2 se Sa eebe eee cee ex T. nigrescens p. 54 1. Not blackening. 2. Gills free, adnexed or sinuate. 3. Pileus acutely umbonate. 4. Gills —1.5 mm wide, very crowded. Smell strong ...................-. T. papillata p. 63 10 TROGIA IN TROPICAL ASIA AND AUSTRALASIA 4° Gillstmuchiwiders subdistant =e soee toe ee ee eee T. tricholomatoides p. 83 3. Not papillate-umbonate. 5. Gills in | rank, distant, adnexed or free. Pileus sulcate, delicate ....... T. delicata p. 26 5. Gills in 3-4 ranks, sinuate, crowded. Pileus firm, not sulcato-striate Sie og bsinlove tebe ov ekewen ald vee tora vat es i MRP aia sea ayoaravane td a eee OPM eee aya T. endocystidiata p. 29 2. Adnato-decurrent to deeply decurrent. 6. Gills 0.5 mm wide. Pileus 2-20 mm wide. 7. Pileus acutely umbonate, —5 mm wide. Gills in 1 rank, often fold-like ROC et eo TCE ICL oe OE: Oke cat og cao DET DRPOIS ROLES Co dco T: mammillata p. 49 7. Not so. 8. Gills in 1 rank, not forked. 9s Spores 4—5:5) pm) widesspalevamylotdie geese reas T. omphalinoides p. 61 (Not amyloid, see 7: octava). 9. Spores 3.5-4.3 x 2-2.3 wm, not amyloid. Smell nitrous ...... T. diminutiva p. 28 8. Gills in 2-3 ranks, often forked. Spores 2.5-3.5 wm wide, not amyloid PE a OE On OAM oa doe ROTC. aD DOO sooo On OEM come Ee T. alba p. 16 6. Gills broader. 10. Pileus 4-9 cm wide, pale yellow, thin. Gills often fold-like, disappearing outwards te ec eee as Cor Ear oe dome On AG ooo Oca nm aaa ook T. macra p. 47 10. Not so. 11. Flesh with copious watery juice. Stem more or less excentric ...... T. seriflua p. 70 11. Not so. 12. Pileus sulcato-striate. 13. Gills 2-6 mm wide in 2 ranks, thick. Stem —25 mm long 7: revoluta p. 68 13. Gills 1-2 mm wide, mostly 1 rank, edge fimbriate. Stem —11 mm long PION ace tea RR Toye TOC cI Ca elie oie lg vib Ae T. polyadelpha p. 64 12. Not sulcato-striate. 14. Gills in 1(—2) ranks. 15. Spores 4.7-6 x 4.5-5 pm, not amyloid .............. T. exigua p. 31 Sin Sylow (G7) SC BIV/ELES) prio, BWC! 7 saeco ocaneoe T. impartita p. 40 15. Spores 6-9 x 4.5-6 wm, not amyloid ........... T. raphanolens p. 67 14. Gills in 2-4 ranks. 16. Base of stem with a fibrillose arachnoid felt. Spores pale amyloid FS REO do OO aN Ole ER SS T. omphalinoides v. confertifolia p. 61 16. No such basal felt. 17. Cheilocystidia as a sterile gill-edge. Spore not amyloid. [82 Pleurocystidias presente sense tsa earn enon T. decipiens p. 25 18. Pleurocystidia absent. 19S Gills lesen Wid elem eretetar ae seen T. limonospora p. 45 19% (Gills)2=4"mmni widew a.) > meena T. marasmioides p. 50 17. Cheilocystidia none or sparse; gill-edge fertile. Spores pale amyloid. 20. Pileus pale yellowish. Spores 5.5-6.5 wm wide T. icterinoides p. 39 20. Pileus white or greyish. Spores 3.5-5 um wide 7: impartita p. 40 KEY TO THE SPECIES ON MICROSCOPIC CHARACTERS il il, 2. Spores amyloid. 3. Fruit-bodies white 3, Fruitsbodiesncoloune dl. wcrc cie« axon cysheus ovara > cher here art eee Group C p. 13 2. Spores not amyloid. 4. Fruit-bodies white 4 BruitsbodiesnCoOlouneds . scij cee «hohe cis cPevtienel ote cr clone larapcl teers erecta erst enone rane eT aeRe Group E p. 14 Pleurocystidia present or, if without gills, then with hymenial cystidia ................. Group A Pleurocystidia absent. en oan Ter ie Nia Ne Bin ote, OO ie ona Ga Group B p. 12 ERAT Sor EE MPa ca 6 Sidi ds cainiion cioo.chaltes oe Group D p. 13 KEY ON MICROSCOPIC CHARACTERS ll Group A 1. Fruit-bodies white, yellowish to pale ochraceous, in one species blackening. Gills well formed but often sterile for some time. Pileus not umbonate, without a pseudoparenchymatous hypodermis. 2. Fruit-body blackening, exuding a watery juice turning red or violaceous, then blackening. Gills adnate, very crowded, narrow. Spores 5-6 x 4-5 um, amyloid ............ T. nigrescens p. 54 2. Not blackening. 3. Flesh of pileus with endocystidia. Spores not amyloid. 4. Gills sinuate, 2-4 mm wide. Cheilocystidia clavate to ventricose fusiform, appendaged. Spores alae Se LITT Th ap ca it ln Oke gr Be ar Sd T. endocystidiata p. 29 4. Gills adnato-decurrent, 1-2 mm wide. Cheilocystidia mostly with a subglobose apex. Spores 6.5-9 3. Without PASS), PITA oe ee, ee ee. = SR EAT Pe Ba T. limonosporoides p. 46 endocystidia. 5. Pileus small, 7-15 mm wide. Stem very short, sublateral. 6. Spores 4 x 2.5 um, amyloid. Caulocystidia with pale brown walls, turning pink in KOH Se ee RR neeeEG Sake as ss eR RSSe eee eee ees os T. sublateralis p. 77 6. Spores 7-9 x 3.5-4.5 um, ? not amyloid. Caulocystidia colourless. Gills —2 mm wide Cee ee Ses sas ote e aeR ewe. Cee eS, T. decipiens v. pleurotella p. 26 5. Mesopodal or somewhat excentric. 7. Pileus shallowly sulcato-striate, —20 mm wide. Stem —10 x 1.5 mm. Secondary gills fold-like. Spores 7-9 x 5.5-6.5 um, amyloid. Basidia 2-spored. Pleurocystidia sparse ee SS ee: 5 Si AO ee SS CURR, 2 es T. polyadelpha p. 64 7. Not sulcato-striate. Secondary gills well developed. Basidia 4-spored. 8. 8. Spores subglobose c. 6 x 5.5 um, amyloid. Tissue with stout oleiferous hyphae 8-20 um wide, ending as hymenial cystidia. Stem usually excentric. Flesh with copious watery UNC ee Stet ana ee eres oh PE Pe cee Ee ete, artes ee T. seriflua p. 70 Spores ellipsoid. Without such oleiferous hyphae. 9. Spores 6-9 x 4.5-5.5 um, not amyloid. Pileus —22 mm wide. Gills —2 mm wide. Plenrocystidiatabundantiz aeons ee ee Oe ee ee es ee T. decipiens p. 25 9. Spores 5.5-7 x 2.7-3.5 um, very faintly amyloid. Pileus —8 mm wide. Gills —0.5 mm wide. Pleurocystidia none but with scattered ends of oleiferous hyphae in the Hy nremaieee. oo tae aa es So. es Se Ee ...2o T. alba v. minor p. 16 1. Pileus coloured or, if pale yellowish, then with a pseudoparenchymatous hypodermis and fold-like gills (7. macra). Spores not amyloid (except T- tenax). 10. Stem short, almost lateral. Wholly cinnamon fawn tinged orange. Pileus —25 mm wide. Spores BA SGne-2- Sigunspale Amyloid 94/5 scs sek sn KOS AS es MS Sn SEG LO T. tenax p. 81 10. More or less mesopodal. Spores larger. 11. Pileus umbonate or, if not, then the gills not decurrent; with a pseudoparenchymatous hypodermis (except 7: subtomentosa). Gill-edge sterile. 12. Pileus acutely umbonate, often rugulose. 13 13. . Gills absent or as slight folds. Spores 7-11 x 4-6 um. A> Pileus. crimsonto'rese jpink(s 245: 32. Bee Sees Se Fak. ss SE T. rosea p. 69 14. Pileus brown, fuscous, ochraceous or greyish ............. T. silvestris p. 73 Gills well formed. 15. Spores globose 8-10 um. Gills orange to deep yellow. Solomon Isl. a TL ASSESS FIEE. IGS, AS OP Be 8 88 T. aurantiphylla p. 18 15. Spores ellipsoid 6-9 x 4-6 um (except T. carminea). 16. Gills decurrent, crowded, 1-2 mm wide, white. Pileus pale fuscous umber. Stemiwhite: 422. - 2. - eee eer ee ees. Sue eeo T. fuscoalba p. 34 (pleurocystidia thick-walled ..... levee 25 rt RGAE var. metuloidea) 16. Gills adnexed, adnate or subdecurrent, usually wider. 17. Pileus yellow with fuscous centre. Stem yellow. Gills white, edge yellow. Pleurocystidia —200 um long with clavate apex 10-17 um wide Shee ards otek terse ens Re eis ms wt. T. fuscolutea p. 36 17. Differently coloured. Gills without yellow edge. 18. Pileus —9 mm wide, rose-red. Gills nearly free. Stem white. Spores Se e235 ps See oe beast ee ae T. carminea p. 21 12 TROGIA IN TROPICAL ASIA AND AUSTRALASIA 18. Larger, differently coloured. 19. Pileus fulvous cinnamon to fawn ochraceous, fuscous or greyish. Stemiewhite’, 2 Acct tavteey- cae enna «a T. fulvochracea p. 31 19. Pileus fuscous umber tinged lilaceous ... T. lilaceogrisea p. 44 12. Pileus not so strongly umbonate. Gills adnexed, adnate or subdecurrent. 20. Pileocystidia none. Caulocystidia —50 x 17 wm, widely clavate. Pileus —3 cm wide, fuscous grey, stem) fuliginous prummoser. -- 4 escmiee ace cre el T. mycenoides p. 53 (smaller ypileusprosesredi= sae eke eee eee oe oe ee T. carminea) 20. Pileocystidia present. Caulocystidia longer, subclavate to ventricose. 21. Pileus subtomentose with septate hyphal ends, fawn ochraceous to fuscous or sepia. Gills ventricose. Spores 5.5-7 x 4.2-5.5 um, slightly amyloid FA AR) ahs Ee OO cei PPR OOS GENES 6S aoc T. subtomentosa p. 79 21. Pileus not subtomentose. 22. Pleurocystidia thick-walled. Pileus fuscous brown. Spores 7-9.5 x 4-5.5 um, notvamyloide Ase. eee: T. fuscoalba v. metuloidea p. 35 22. Pleurocystidia thin-walled. Smell often farinaceous. 23. Pileocystidia —250 um long, with supporting hyphae at the base. Pileus —3 cm wide, fuscous umber to fuscous ochraceous. Stem 1-3 mm thick + ageettiencatnGGue vie tens Peay ers captors ope ap vars ees... T. hispidula p. 38 23. Pileocystidia shorter, not so supported. Pileus —9 cm wide, fuscous vinaceous to greyish lilac. Stem 2-7 mm thick. Smell strong weld . Len wemeshe 206, adstere CR « SNe T. lilaceogrisea p. 44 11. Pileus infundibuliform, not umbonate. Gills usually deeply decurrent. 24. Wholly pale yellowish. Pileus 3-9 cm wide, thin. Gills 0.5-1.5 mm wide, often fold-like, edge fertilesisporessd—l 0-425 Oy ae ieee eee T. macra p. 47 24. Not so. Gill-edge with cheilocystidia. Pileus fuscous, greyish brownish or greenish. 25. Pileus without a pseudoparenchymatous hypodermis. Spores ellipsoid. 26. Pileus —10 mm wide. Pleurocystidia scarce. Cheilocystidia mostly clavate as a Sterileyedgem Spores!6=765) >e4—Siym eee eee T. calyculus p. 21 26. Larger. Cheilocystidia ventricose lanceolate, not as a sterile edge. Spores larger. 27. Pileus grey green. Gills —3 mm wide. Spores 7-10 x 4-5 um, not amyloid 5.2L AD GRRL EAS ee ee ee eee, BE T. subviridis p. 80 27. Pileus fuscous. Gills 0.5 mm wide, often forked. Spores 8-10 x 6.5-8.5 ym, palecamyloidwan. C292 Se ee NP T. furcata p. 33 25. Pileus with a pseudoparenchymatous hypodermis, not greenish. 28. Spores 8-10 x 6-7 um, ellipsoid. Cheilocystidia more or less ventricose. Flesh Waxy-soft, tough) therpills/ secre eee ele ree eee T. cystidiata p. 25 28. Spores 6-8 x 5.2-7 wm, subglobose. Cheilocystidia mostly clavate. Flesh tough, subgelatinous. Lignicolous or humicolous ........... T. subgelatinosa p. 75 Group B 1. Gills none or as slight folds not reaching the stem. Hymenium thickening. Cystidia none. Pileus —]5 mimi Wides Spores: d—8:9) e429 —O 9 Mile seyeeeeee) eevee te eeade sesiarais ini eestepeenes