Karstenia 28:65-77. 1988 (1989) The Tricholoma caligatum group in Europe and North Africa ILKKA KYTOVUORI KYTOVUORI, I. 1989: The Tricholoma caligaturn group in Europe and North Africa. - Karstenia 28:65-77. Three species are recognized in the Tricholoma caligatum group in Europe: T. caligatum (Viv.) Ricken, T. nauseosum (Blytt) Kytiiv., comb. nova, and T. dulciolens Kytov., spec. nova. The first two species are also reported from North Africa. The East Asian T. matsutake (S. Ito & Imai) Singer is included in T. nauseosum, which is thus an edible fungus of major international interest. The distribution of each species in the study area is mapped, and their ecology and relationships are discussed. llkka Kytovuori, Department of Botany, University of Helsinki, Unioninkatu 44, SF-00170 Helsinki, Finland In Europe (e.g. Moser 1983) the Tricholoma caliga­ rium of Soc. Catal. Micol. Barcelona. Types and tum group is traditionally supposed to be represented some other specimens were also examined from by one species only. One species is also recognized MICH. in East Asia, whereas in North America Smith (1979) Microscopical characters were examined with the reports four species and four varieties of the group. microscope Leiz Laborlux 12 at magnifications of However, heterogeneity is evident in the group in about 625 and 1 560. Spores, basidia and surface hy­ Europe as well, though to a smaller extent than in phae were drawn with the Leiz drawing tube at a North America. Smith (1979) and most other North magnification of 2 OOO. Spores were examined, mea­ American authors, e.g. Pomerleau (1980), refer the sured (with an ocular micrometer) and drawn from T. caligatum group to the genus Armillaria. In this slides showing the surface view of pieces of gills paper I follow the treatment of most European au­ and/or hyphae from the ring or stipe top of dried ba­ thors, e.g. Singer (1986), Moser (1983), Bon (1976, sidiocarps, mounted in Melzer's reagent. Only spores 1984) and Kost (1981), and refer the group to the in a horizontal position were measured; young, genus Tricholoma. anomalous, very large and gigantic spores were ex­ cluded. When possible, both sclerospores and hya­ line spores were measured. In some specimens all the Material and methods hyaline spores were somewhat collapsed and then only sclerospores were measured. Length and width Specimens were checked in the following herbaria were measured from the same spore and the length/ (all in Europe): AQUI, ANGUC, B, BCC, BR, width ratios were calculated for individual spores. BRA, BP, BREM, BRNM, C, DBN, FI, G, GB, H, Basidia were drawn and measured from the same HBG, HOH, IB, JE, Joensuu Research Station, K, slides as the spores. Both sclerobasidia and hyaline KUO, L, LG, LISU, LY, M, MA, MPU, 0, OULU, basidia were drawn, but most of the measurements PC, PAD, PAV, PERU, PRM, S, TRH, TUR, were made from sclerobasidia because the hyaline Univ. of Joensuu, UPS, VER, W (the symbols ac­ ones were often somewhat collapsed. Cap and stipe cording to Holmgren et al. 1981). Some specimens surfaces were examined from tangential sections were examined from the private herbaria of Dr. M. made with a razor blade. Bon, Saint Valery-sur-Somme, Dr. D. Lamoure, The illustrations are my own unless otherwise Lyon, and Dr. A. Riva, Balerna, and from the herba- stated. 66 KYTOVUORI: TRICHOLOMA CALIGA TUM GROUP KARSTENIA 28 (1988) Key to species broadly ellipsoid, hyaline, smooth, apiculus distinct. Some sclerospores present, with thickened, hyaline Odour strong, sweetish, like Inocybe corydalina, stipe an- to somewhat dextrinoid wall. Sclerospores somewhat nulate .................................. ............................... 2 smaller and relatively wider than the normal spores. - Odour different (e.g. farinaceous or of coal gas) or lacking, Spore contents hyaline, appearing very indistinctly stipe annulate or not . ... ...................... Tricholoma spp. 2 Cap small (to 10 cm), fuscous to blackish, ± Mediterranean coagulated to guttulose in Melzer's reagent. species ............................................. I. T. caligatum Basidia clavate, 27-42 x 5.5-7.5 µm (149 basidia - Cap small to very large (5 to 25 cm),± brownish, with or of 28 specimens), 4-spored, with fairly large sterig­ without cameous tint, temperate toboreal species ... .... 3 mata. Sclerobasidia sparse, with thick, most often 3 Cap large, robust (to 25 cm), with large, appressed scales, hyaline, rarely dextrinoid wall. No differentiated cys­ without cameous tint, spores more than 6.5 µm long, with Pin us ............................................. 2. T. nauseosum tidia, but some clavate free ends of hyphae present at - Cap smaller (to 10 cm), with numerous very thin, small the gill edge. scales, with cameous tint, spores less than 6 µm long, Pileal surface formed by more or less flat hyphae with Picea ....................................... 3. T. dulciolens 7-16 µm wide, dark brown to fuscous in Melzer's reagent. The uppermost hyphae somewhat lighter in colour, flatter and thinner-walled than the lower ones. Pigment evenly distributed, no incrustations present. 1. Tricholoma caligatum (Viv.) Ricken, Stipe surface below the ring like that of the pileus. Figs. 1-3, 6. Above the ring tufts of torulose and forked entangled Tricholoma caligatum (Viv.) Ricken, Blatlerpilze Deutsch!. hyphae and large, clavate hyphal ends with hyaline 331. 1915. - Agaricus caligatus Viv., Fungh. ltal. 40, pl. but distinct wall. 35. 1834. Armil/aria caligata (Viv.) Gillet, Hymen. France. 79. 1874. - Type: not typified. Armillaria caligata f. gracilis Bres., Icon. Mycol. l. pl. 45. 1927. - Type: not seen. Distribution and ecology Illustrations. Viviani (1834 pl. 35), Baria (1888 pl. 18), Tricholoma caligatwn seems to be a meridional spe­ Bresadola (1927 pl. 45), Dahncke & Dahncke (1980 p. 155), cies occurring most commonly around the western Bon (1976 fig. 66, 1984 fig. 66, 1987 p. 161), Soc. Cata!. Micol. (1985 pl. 197), Marchand (1986: 895), Moser & Jiilich part of the Mediterranean, especially in South France, (1987 pl. III: 1), Lanzoni (1988). - Not Cetto (1980 pl. 603), the eastern and southeastern parts of Spain and Pilat & U� (1961 figs. 62, 63), Michael et al. (1979 fig. adjacent NW Africa (Maire 1915, Maire & Werner 204, 1987 fig. 204), Korhonen (1986 p. 144), Riva (1988 pl. 1937, Malern;on & Bertault 1975, Bon 1976, 1984, 68). Moreno 1986). With one exception, all the rather few Pileus 3-12 cm wide, subumbonate to plano­ specimens of the present group collected from France convex; margin inrolled when young, outrolling with and Spain and seen by me belong to T. caligatum age. Surface weakly viscid when wet, somewhat s.str., and thus the literature records of T. caligatum shining when dry, smooth and blackish brown to from that region are best referred to that species. fuscous in the centre, with appressed, large, dark According to Bon (1976, 1984), in France T. cali­ brown scales towards the margin, at the margin often gatum grows in acid Pinus forests, but also with chapped, exposing the pale pileal trama, under the Abies, Picea and Quercus suber or other evergreen lens radially striate, not costate. Context thick in the Quercus species in macchia. According to the speci­ centre, thin towards the margin, whitish, and fairly men labels, the most important tree species is Pinus compact. Odour strong, just like that of Inocybe halepensis, the other species mentioned being P. pi­ corydalina or Pholiota heteroclita. Taste sweetish-bit­ nea and Quercus ilex. Bon (1976) also reports the terish to bitter. species under Picea and Abies at higher levels in the Lamellae close, broad, straight, sinuate, whitish, French Central Upland, but the identity of the species somewhat anastomosing, with some lamellulae. there needs to be confirmed with specimens. The Stipe 4-10 cm long, 1-2.5 cm wide, with an as­ only specimen from there seen by me (Puy-de-Dome, cending persistent annulus 7-25 mm down from the Lagoutelle, Herb. Bon 70161) was T. nauseoswn. T. lamellae. Surface below the ring concolorous with the caligatwn s.str. from Pralognan, 1 450 m (Herb. pileus, with fairly wide, more or less transverse, Lamoure), represents the highest record of the spe­ blackish brown zones on a lighter background, above cies. The northeasternmost record of T. caligatum the ring whitish. Outer side of the ring base of the s.str. is from Baden-Wiirttemberg, West Germany colour of the stipe below, the inner side and the edge (HOH, OULU). Two of the specimens from the latter white. Context whitish, compact, solid. area were collected on calcareous soil. Some Spores 5.7-7.3 x 4.3-5.4(-5.9) µm (431 spores heterogeneity may still exist in T. caligatwn in Central of 34 specimens), length/width ratio 1.20-1.40, and South Europe. KARSTENIA 28 (1988) KYTOVUORI: TRICHOLOMA CALIGATUM GROUP 67 Tricholoma caligatum 240 480 Fig. 1. Distribution of Tricholoma caligatum in Europe and adjacent North Africa according to the material examined. Asterisk indicates the type locality. From NW Africa I have seen T. caligatwn s.str. seems to be from late October to early December, but gathered in Pinus halepensis forests only. All the in the warmest regions basidiocarps may possibly de­ specimens from Cedrus atlantica forests were T. nau­ velop throughout the winter. seosum. Viviani (1834) described T. caligatum from Liguria in Italy. From there the distribution stretches southward. The southeastemmost record is from Cy­ Discussion prus. The fact that Bresadola (1827) described the present taxon as forma gracilis possibly indicates that The nomenclature of the T. caligatwn group in Eu­ it is the rarer of the two species that occurs in North rope is unclear, apparently owing to confusion with Italy. the T. robustum-T. focale complex. Viviani's (1834) The fruiting period of T. caligatwn s.str. is long description of T. caligatwn is accurate and Barla and it occurs late in the year (Fig.