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Mycologia, 90(3), 1998 pp. 494–504. © 1998 by The New York Botanical Garden, Bronx, NY 10458-5126 The pantropical genus Macrocybe gen. nov. David N. Pegler1 grande in section Leucorigida. Both the type species The Herbarium, Royal Botanic Gardens, Kew, of this section, T. mongolicum S. Imai, and the second Richmond, Surrey TW9 3AB, UK cited species, T. altaicum Singer, are common species in northern China and Mongolia, where they form D. Jean Lodge2,3 fairy rings in grass. They are remarkably similar to Center for Forest Mycology Research, Forest Products the North temperate species, Calocybe gambosa (Fr.) Laboratory, USDA-Forest Service, Luquillo, Puerto Rico 00773-1377 Donk and its allies, and must be regarded as belong- ing to Calocybe Kühner ex Donk. 3 Karen K. Nakasone The remaining species form large, robust basid- Center for Forest Mycology Research, Forest Products iomata that are tricholomatoid in habit, but their pre- Laboratory, One Gifford Pinchot Drive, Madison, cise position within the genus nevertheless remains Wisconsin 53705-2398 problematical. In many ways, these species closely re- semble those of Calocybe, in tribe Lyophylleae Kühner ex Bas. However, siderophilous granules are absent Abstract: Analysis of morphological and ecological from the basidia of fresh and dried specimens (C. characters plus ribosomal DNA sequences support the removal of seven related tropical species from the Ovrebo, pers. comm.; Pegler, 1983, respectively), and genus Tricholoma. This group of apparently nonmy- the presence of these granules characterizes tribe corrhizal species that have abundant clamp connec- Lyophylleae. Siderophilous granules may be weakly tions on all hyphae but lack siderophilous granula- stained in the basidiospores. However, the presence tion in the basidia cannot be accommodated in other of clamp connections throughout the basidiomata, genera, such as Calocybe or Megatricholoma. We there- cyanophilic spores, and the nonectomycorrhizal na- fore propose Macrocybe as a new genus in the Tricho- ture, provide a unique combination of characters. lomataceae for these taxa. Kost (1984) proposed the genus Megatricholoma to Key Words: Agaricales, Calocybe, ribosomal DNA, accommodate the European species, Tricholoma co- Tricholoma, Tricholomataceae, Tricholomateae lossum (Fr.) Quél. This species forms large basidiom- ata but differs from both Tricholoma (Fr.) Staude and the tropical species in its hemiangiocarpic develop- ment resulting in an annulate stipe, together with INTRODUCTION differences in the subhymenial layer and basidia, and Singer (1986) included only one genus, Tricholoma the presence of true cheilocystidia and pleurocysti- (Fr.) Staude, when he proposed the subtribe Tricho- dia. Furthermore, M. colossum lacks clamp connec- lomatinae. This is characterized by basidiomata that tions. form obligate ectomycorrhizal associations with for- The striking appearance of these large, pale, non- est trees, and hyphae that lack clamp connections. ectomycorrhizal, tropical and subtropical agarics, Exceptions to this definition are found in subgenus which are distinct from Calocybe, Megatricholoma and Contextocutis Singer, Section Leucorigida Singer, as Tricholoma sect. Leucorigida sensu stricto, require rec- proposed by Singer (1945). Singer (1986) included ognition at the generic level. We present a molecular T. titans (as T. cystidiosum), T. giganteum and T. prae- analysis by Nakasone and Lodge of the 5'-end of the ribosomal large subunit DNA sequences of ‘T’. titans, Accepted for publication November 17, 1997. four ectomycorrhizal species of Tricholoma (T cf. fla- 1 Email: [email protected] vovirens, T. imbricatum, T. cf. intermedium, and T. vac- 2Email: [email protected] cinum), Clitocybe clavipes and Ca. gambosa. The mor- 3This article was written and prepared by U.S. government em- ployees on official time, and it is therefore in the public domain phological and ecological data, considered together and not subject to copyright. The Forest Products Laboratory in with molecular analyses by us and others are consis- Madison is maintained in cooperation with the University of Wis- tent with our hypothesis that T. titans falls outside consin, while the one in Puerto Rico is maintained in cooperation with the International Institute of Tropical Forestry, USDA-Forest the Tricholoma sensu stricto clade, and placement of Service. T. titans in the genus Calocybe of Tribe Lyophylleae is 494 PEGLER ET AL.: MACROCYBE 495 also not strongly supported. A new genus, Macrocybe, (1975). Spore dimentions, means, standard devia- is therefore proposed. tions and Q’s are given for 25 spores taken from the Some of the tropical species belonging to the pro- type or an authentic collection, except for two col- posed new genus have had a complex and confusing lections that were sequenced and are described be- history. Petch (1912: 268) supplied a revised account low. of T. crassum (Berk.) Sacc., based on material col- lected in the Kandy District of Central Province, Sri The following taxa were included in the molecular study: Lanka. At that time, he placed another species that T. titans (Lodge PR-1637, see specimens examined for data; was also originally described from Sri Lanka, T. pa- GenBank U8637-U8638), T. cf. flavovirens (Pers. : Fr.) Lun- dell (USA, WISCONSIN: Sauk County, County Highway PF, chymeres (Berk. & Broome) Sacc., in synonymy. This in Picea plantation, Oct. 1994, T. J. Volk TJV 94-134, CFMR, caused confusion between the two species that has GenBank U86672), T. vaccinum (Pers. : Fr.) Kummer (USA, persisted until now. Tricholoma pachymeres has been WISCONSIN: Oct. 1994, T. J Volk TJV 94-133, CFMR, recorded on a number of occasions (Baker and Dale, GenBank U86443-U86444), Ca. gambosa (Fr. : Fr.) Singer 1951: 94, pl. 1g; Pegler, 1983: 111, pl. 3D), and was ex Donk (ENGLAND, SURREY Esher, West End Common, regarded as a pantropical species. Cifuentes and Guz- copse under young Ulmus and Sambucus, B. M. Spooner, man (1981) described a similar species from Mexico K(M)22752; GenBank U86440-U86441), Cl. clavipes (Pers. : as T. cystidiosum Cifuentes & Guzmán, which presum- Fr.) Kummer (USA, WISCONSIN: Sauk County, Oct. 1994, ably differed in the presence of hymenial cystidia. T. J. Volk TJV 94-122, CFMR, GenBank U86439-86440). The However, the type material of the paleotropical spe- following sequences obtained from GenBank were also in- cies, T. pachymeres, is in poor condition so it is not cluded in the analysis: T. cf. intermedium Pk. (USA, CALI- possible to confirm the presence of pleurocystidia FORNIA: Sierra County, Tahoe National Forest, Rd. 54 off Highway 49, near Yuba Pass, buried groups under old- and hence whether or not it is conspecific with T. growth Abies magnifica, 15 Oct. 1995, N. Wilson, ledger titans. Courtecuisse renamed the neotropical species K.M. Shanks KMS 593, SFSU; GenBank U76465), T. imbri- as T. cifuentesii in 1985 to avoid confusion with an catum (USA, CALIFORNIA: Marin County, Pt. Reyes, 9 Jan. earlier species, T. cystidiosum A. H. Sm. Singer (1990) 1993, K.M. Shanks KMS 356, SFSU; U76458), Pleurotus pul- synonymized T. cystidiosum Cifuentes & Guzmán with monarius (Fr. : Fr.) Quél. (U04153) and P. cornucopiae (Pau- T. titans H. E. Bigelow & Kimbr., a species described let) Rolland (U04135). The sequences for T. intermedium one year earlier from Florida (Bigelow and Kim- and T. imbricatum were provided by K. M. Shanks before brough, 1980), again emphasizing the presence of submission to GenBank. hymenial cystidia. Recent Puerto Rican collections by one of us (DJL) The culture of Cl. clavipes and the specimens of match the characteristics of T. titans well. The char- Cl. clavipes, T. vaccinum and T. cf. flavovirens are acters of T. titans are distinct from those of T. gigan- deposited at CFMR. The specimen of Ca. gambosa is teum Massee from West Bengal, India, and T. prae- at K, and the specimens of T. intermedium and T. grande from Brazil. The latter two species lack the vaccinum are deposited at SFSU. The specimens were squarrose appearance of the stipe associated with the typical for their species, except for T. cf. flavovirens numerous small, recurved squamules as found in T. and T. cf. intermedium, which are briefly described titans, T. crassum and T. pachymeres, and they also below. lack pseudocystidia. A reexamination of all the neo- Extraction of total DNA.—Total DNA was obtained tropical collections deposited at Kew under the name from basidiospore drops, dried basidiomata, or ly- T. pachymeres’ has demonstrated the presence of ophilized cultures. For T. vaccinium and M. titans, a pseudocystidia and other characteristics consistent basidiospore suspension in 50 ml of TE (10 mM Tris, with T. titans. An examination of all the collections 1 mM EDTA, pH 8) was placed in an 1.8-ml micro- from Costa Rica deposited at the Field Museum of Natural History under the name ‘T. praegrande’ also centrifuge tube with a small amount of glass beads demonstrated the presence of cystidia and scaly stipes (Sigma G-4649), vortexed at high speed for 2 min, consistent with T. titans. In addition to describing a and stored at -20 C. new genus to accommodate these tropical and sub- Dried lamellae or small sections (1 X 1 cm) of the tropical species, we provide the correct placement of pileus of T. cf. flavovirens and Ca. gambosa were various collections under the material examined sec- ground to a fine powder with a small amount of sand tion for each species. with a mortar and pestle, and DNA was extracted fol- lowing Cenis (1992). In the case of Cl. clavipes, a culture was grown on 50 mL of sterile 2% malt ex- MATERIALS AND METHODS tract supplemented with 1% glucose at 25 C for 1 wk. Color names in quotation marks are those in Ridgway The culture was harvested by filtration onto Mira- (1912) or Ridgway colors as reproduced by Smithe cloth (Chicopee Mills, Inc., La Jolla, California), ly- 496 MYCOLOGIA ophilized, and stored at -20 C in plastic bags. Total tomycorrhizoidea. Species typicae: Tricholoma titans DNA was extracted following Cenis (1992).
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