Leccinum (Boletaceae) in Costa Rica

Transcription

1 Mycologia, 95(3), 2003, pp by The Mycological Society of America, Lawrence, KS Leccinum (Boletaceae) in Costa Rica Roy E. Halling 1 Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York Gregory M. Mueller Department of Botany, The Field Museum of Natural History, Chicago, Illinois Abstract: Complete descriptions, illustrations, distributions and a key to species are provided for the eight known Costa Rican species of Leccinum. Leccinum tablense is newly described from oak forests, and Leccinum monticola is newly described from the páramo and adjacent timberline oak forests where it appears to be associated with Comarostaphylis arbutoides a member of the Ericaceae. All are compared to similar taxa. Key words: biogeography, boletes, Central America, Comarostaphylis, ectomycorrhizae, oaks, Quercus INTRODUCTION The mycelium of Leccinum is directly connected with ectomycorrhizal tree roots, according to Singer (1986). Halling (1999) recently stated that species of Leccinum are uncommon in the neotropics, compared with northern temperate forests, due primarily to the paucity of suitable phanerogamic associates (such as Pinaceae, Betula, Quercus and Populus). For example, south of Nicaragua, Quercus appears to be the only traditional associate where it forms canopy stands at montane elevations (generally above 1500 m) in the Cordillera Talamanca. Studies by Kapelle et al (1992) and Kapelle (1996) have provided extensive data on the phytogeography and phytosociology of Talamanca Quercus forests in Costa Rica. Halling (1989) described the first neotropical Leccinum (L. andinum) from Colombia and summarized the work on South American Boletaceae completed at that time. One of us (Halling 1999), described two additional new species of Leccinum with a summary listing of five others known from Costa Rica. One of those, L. quercinum Pilát, was misidentified but, based on our recent observations, we describe it below as L. Accepted for publication December 12, Corresponding author. rhalling@nybg.org monticola, and we believe it forms ectomycorrhizae with Comarostaphylis, not Quercus as previously thought. With regard to the mycorrhizal status of Comarostaphylis, Bidartondo and Bruns (2001) published a phylogenetic tree of some Ericaceae (Monotropoideae in particular) with one clade containing Comarostaphylis arbutoides Lindley, Arbutus menziesii Pursh., Arctostaphylos manzanita Parry, A. uva-ursi (L.) Spreng., Pyrola picta Sm. and P. aphylla Sm. There was a notation next to those six taxa stating they form mycorrhizal associations with diverse Ascomycetes and Basidiomycetes. However, documentation justifying that statement was lacking. Subsequently, M. Bidartondo (pers comm) showed the senior author digital images of arbutoid roots from the plant they sampled (origin: Chiapas, Mexico) growing as an exotic in the University of California at Berkeley Botanical Garden. We would submit that this paper is a first report of such a putative relationship because L. monticola appears solely with plants of C. arbutoides above timberline in the páramo and just below timberline, where there is a mixture of that ericad and Q. costaricensis Liebm. Considering that the native distribution of C. arbutoides is central Chiapas, Mexico, through Guatemala, Honduras, Nicaragua, Costa Rica and western Panama at elevations of m ( J. Luteyn pers comm), the senior author is in the process of determining the mycorrhizal status of L. monticola and C. arbutoides. Prior documentation of Costa Rican Boletaceae, as well as a hypothesis and documentation supporting co-migration of Quercus and ectomycorrhizal fungi to the montane neotropics, were summarized by Halling (1999) and Halling and Mueller (1999, 2002). Circumscription and subgeneric placement of taxa are according to Singer (1996) and Lannoy and Estades (1995). MATERIALS AND METHODS The descriptive data given here were produced from a DEL- TA database (Dallwitz 1980, Dallwitz et al 1993 onward). That database also contains an interactive identification and information retrieval system using the program Intkey, running under MS-Windows (Dallwitz et al 1998) and is accessible on the Internet in both English and Spanish at: Color designations (e.g., 4A3) in the species descriptions are from 488

2 HALLING AND MUELLER: COSTA RICAN LECCINUM 489 Kornerup and Wanscher (1983). FIGURES 1 8 were produced digitally, according to the protocol outlined by Huhndorf at collections/ botany/botany sites/imagemanage/intropage.htm. Microscopic features were drawn with a drawing tube affixed to an Olympus BHS-2 compound microscope equipped with Nomarski DIC optics. Q mean length/width ratio for 20 spores measured. Herbarium acronyms are from Holmgren et al (1990). TAXONOMY KEY TO LECCINUM IN COSTA RICA 1. Hymenophore yellow, bluing when injured... L. andinum 1. Hymenophore usually not yellow, never bluing when injured Pileus rugulose during some stage of development; pileipellis hymeniform Pileus even throughout development; pileipellis not hymeniform Context becoming pink to reddish orange in the pileus and upper part of the stipe when exposed Context unchanging Context not changing to blue-green or dark blue in the stipe base when exposed; stipe scabers caramel to light brown...l. rugosiceps 4. Context changing to blue-green to dark blue in the stipe base when exposed; stipe scabers dark gray to black..... L. talamancae 5. Pileus dark reddish brown; hymenophore yellow; stipe scabers tan to gray brown; stipe surface developing some red or yellow tones...l. neotropicalis 5. Pileus light gray brown to tan; hymenophore white to orangish white; stipe scabers gray to black; stipe surface white... L. tablense 6. Stipe with fine pink scabers on white ground color over upper part, chrome yellow at the base...l. cartagoense 6. Stipe with dark-colored scabers (brown to black), never chrome yellow in the base Pileus orange brown to red brown, with sterile flap at margin; stipe white with brown to black scabers......leccinum monticola 7. Pileus violet brown to chocolate brown, lacking sterile flaps; stipe pale violet with brown scabers... L. eximium Leccinum andinum Halling, Mycotaxon 34: FIGS. 1, 9 11 Pileus (4 ) cm broad, at first convex, with age plane, subviscid to tacky, entire to subrugulose, becoming even; disk even, at first brownish yellow (5C6), then raw sienna (6D7); margin subrugulose to uneven, when young grayish yellow, then brass yellow (4C7); surface subtomentose, becoming glabrous; maculose color spots present, amber yellow (4B6) to brownish yellow (5C6) to cinnamon brown (6D6) to honey yellow (4-5D6) to light orange (5A5) to yellowish brown (5E8); sterile appendiculate flaps absent (but occasionally with a sterile zone). Flesh 7 20 mm thick, orange white (5A2) to white to yellowish white (3-4A2), staining slightly blue to blue-green; odor mild; taste mild. Hymenophore tubulose, depressed. Tubes 3 20 mm long, light yellow (2-3A6-5), becoming olive yellow (3D6), blue when injured; pores 2 3 per mm, deep yellow to yellow (3A7-4A8), then grayish yellow (2B5), blue-green when bruised. Stipe 3 12 cm long, mm wide, equal to subequal to subclavate, dry, densely scabrous, pale yellow (2-3A6-5) at first, with age grayish yellow (2B5) and scabrous to subglabrous; scabers when young yellow, with age brownish orange to brownish yellow to light brown to yellow ochre; base white, staining not present. Stipe interior solid; flesh above yellow, staining blue; flesh at base marbled brown and white. Basidiospores olive brown to umber brown, m m, Q 2.88, smooth, ellipsoid to subfusoid to fusoid, dextrinoid, with KOH ochraceous. Basidia m m, clavate, hyaline, 4-sterigmate. Hymenial cystidia m 5 11 m, moderately abundant (more common near pores), thin-walled, colored homogeneous contents or hyaline (rare), lageniform to ampullaceous to subfusoid to ventricose rostrate. Tube trama boletoid, hyaline, in KOH yellowish, lateral strata elements 2 7 m wide, subgelatinous. Pileipellis hyphae a trichodermium, in KOH yellow ochraceous (sometimes with ochraceous homogenous contents), inamyloid; elements m wide, cystidioid to inflated to obtuse to slightly tapered to subcapitate, smooth. Intercalary cells subisodiametric, m m. Pileus trama interwoven, hyaline, inamyloid, with elements m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to versiform caulocystidia, m 3 12 m, filamentous with short-branched apices or subcylindric. Clamp connections absent. Habit, habitat and distribution: solitary or scattered to gregarious in soil under Quercus costaricensis, Q. seemannii Liebm. (in Costa Rica), Q. humboldtii HBK (in Colombia). Material examined: Costa Rica. San José: Dota, San Gerardo, 5 km SW of Cerro de la Muerte, Hotel de la Montaña, Savegre, N, W, 2500 m, 8 Jun 1994, Halling 7246 (NY, USJ); 9 Jun 1994, Halling 7250 (NY, USJ); 10 Jun 1996, Halling 7657 (NY, USJ); 11 Jun 1996, Halling 7665 (NY, USJ); 8 Jun 1997, Halling 7705 (NY, USJ); 26 Jun 2000, Halling 7935 (NY, USJ); 3 Jul 2000, Halling 7982 (NY, USJ); 9 Jun 2001, Halling 8151 (NY, USJ); 27 Jun 2001, Halling 8200 (NY, USJ); 5 Jul 2001, Halling 8249 (NY, USJ). San José: Pérez Zeledón, Villa Mills, C.A.T.I.E. Experimental Forest, N, W, 2850 m, 8 Nov 1993, Halling 7145 (NY, USJ); 22 Jun 1994, Halling 7331 (NY, USJ); 9 Jun 1996, Halling 7649 (NY, USJ); 10 Jun 1997, Halling 7715 (NY, USJ); 27 Jun 2000, Halling 7945 (NY, USJ); 30 Jun 2001, Halling 8229 (NY, USJ). The yellow to yellow-brown colors and blue to bluegreen bruising reaction are good field characters. The association with oak, the yellow hymenophore

3 490 MYCOLOGIA FIGS Habits of Costa Rican Leccinum species. 1. L. andinum (Halling 8200). 2. L. cartagoense (Halling 7394). 3. L. eximium (Halling 7798). 4. L. monticola (Halling 8288). 5. L. neotropicalis (Halling 8218). 6. L. rugosiceps (Halling 8163). 7. L. tablense (Halling 8136). 8. L. talamancae (Halling 8168). 1, , 3, 7, FIG. 2 courtesy CABI Publishing.

4 HALLING AND MUELLER: COSTA RICAN LECCINUM 491 FIGS L. andinum (Halling 5052). 9. Caulocystidia. 10. Hymenial cystidia. 11. Basidiospores. Courtesy Mycotaxon. and yellow context indicate placement in section Luteoscabra, but consistent bluing reactions have not been reported previously. First described from montane Colombia near Medellín (Halling 1989), it is known in Costa Rica from sites near San Gerardo and Villa Mills in the Cordillera Talamanca. Leccinum cartagoense (Wolfe and Bougher) Halling and G. M. Mueller in Halling, Kew Bull 54: FIGS. 2, Tylopilus cartagoensis Wolfe and Bougher, Austral Syst Bot 6: Pileus cm broad, at first convex, with age plano-convex, dry to moist; disk even, at first camel brown (6D4) to cinnamon brown (6D6) to cocoa brown (6E4-6) to gray brown (6-7E5), then tan (6C5-4); margin even, usually with pink overtones; surface matted tomentose to subtomentose to tomentose; maculose color spots present, when young rosy pink; sterile appendiculate flaps absent. Flesh 6 10 mm thick, white with thin dark pink zone under pileus surface, staining absent; odor mild; taste mild. Hymenophore tubulose, depressed to deeply depressed. Tubes 7 10 mm long, yellowish white (4A2), becoming grayish brown to pale pink, pale brown when injured; pores up to 1 mm wide, flesh pink (6A-B3) or white, unchanging when injured or pale brown when bruised. Stipe cm long, 6 15 mm wide, equal, dry; upper half finely scabrous, white; lower half scabrous to subscabrous ridged, white; scabers when young rosy pink, with age dull pink to gray; base chrome yellow to cadmium orange (3,4,5A8), staining not present. Stipe interior solid; flesh above white, staining blue (very rarely); flesh at base cadmium orange to chrome yellow (3-4A8), staining not present. Basal mycelium yellow. Basidiospores pinkish brown, m m, Q 2.58, smooth, subfusoid to fusoid to ellipsoid, inamyloid, with KOH golden yellow. Basidia m m, clavate, hyaline, 4-sterigmate. Hymenial cystidia m 8 11 m, more common toward edge of tubes, thin-walled, with goldencolored contents, fusoid to ventricose, gelatinization absent, encrusting pigment absent. Tube trama boletoid, hyaline, lateral strata elements m wide, not gelatinized. Pileipellis hyphae a trichodermium and interwoven, in KOH brownish, inamyloid; elements m wide, cylindric to subelongate to elongated, smooth, thin-walled, granular content absent, not gelatinized. Pileus trama interwoven, hyaline, inamyloid, with elements 5 12 m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, m 8 10 m wide, clavate to ventricose or lageniform (often secondarily septate and with a long mucro), with golden-yellow contents or hyaline, with encrusting pigment absent. Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: solitary or gregarious on soil under Quercus copeyensis C. H. Müll., Q. costaricensis, Q. seemannii, Q. rapurahuensis Pittier ex Trel. in the Cordilleras Central and Talamanca. Material examined: Costa Rica. Alajuela: Grecia, Grecia, Bosque del Niño, N, W, 1900 m, 18 May 1996, Halling 7533 (NY, USJ); 31 May 1996, Halling 7592 (NY, USJ); 15 Jun 1996, Halling 7689 (NY, USJ). Cartago: El Guarco, Parque Nacional Tapanti, Macizo de la Muerte, Area de Conservacion La Amistad Pacifico, N, W; 2600 m, 6 Jul 2001, Halling 8258 (NY, USJ). San José: Dota, La Chonta, S of Interamerican Highway near Cerro Chonta, N, W, 2400 m, 4 Nov 1993, Halling 7123 (NY, USJ). Copey, 12 km S of Copey on road to Providencia, N, W, 2800 m, 20 Oct 1994, Halling 7394 (NY, USJ). Jardín, 3.5 km W of Empalme, N, W, 2220 m, 7 Jun 1997, Halling 7702 (NY, USJ). San Gerardo, Hotel de la Montaña, Savegre, 5 kmsw of Cerro de la Muerte, N, W, 2500 m, 6 Nov 1993, Halling 7133 (NY, USJ); 20 Jun 1994, Halling 7309 (NY, USJ); 21 Jun 1994, Halling 7324 (NY, USJ); 16 Oct 1994, Halling 7365 (NY, USJ); 19 Oct 1994, Halling 7386 (NY, USJ); 21 Jun 1995, Halling 7441 (NY, USJ); 3 Jul 2000, Halling 7970 (NY, USJ); 27 Jun 2001, Halling 8203 (NY, USJ); 5 Jul 2001, Halling 8254 (NY, USJ). First described from one collection on the western slope of Volcan Irazu (Wolfe and Bougher 1993), Leccinum cartagoense is encountered routinely in the Cordillera Talamanca and on Volcan Poas in the Cordillera Central. The fine pink scabers on the stipe

5 492 MYCOLOGIA FIGS L. cartagoense. 12. Caulocystidia (Halling 8254). 13. Basidiospores. 14. Hymenial cystidia (Halling 8258). surface and the intense chrome yellow stipe base are diagnostic. Originally described as a Tylopilus in subgenus Roseoscabra because of the pinkish flesh-colored spore deposit and its resemblance to Tylopilus chromapes (Frost) Smith and Thiers of the Northern Hemisphere, the Costa Rican entity lacks the intense pink of the pileus and is smaller in stature. There is a rare entity of this group in Costa Rica that has a black pileus and seems to lack pink scabers on the stipe. Additional material is needed to assess the autonomy of this latter form. Leccinum eximium (Pk.) Singer, Persoonia 7: FIGS. 3, Boletus robustus Frost non Fr., Bull Buffalo Soc Nat Sci 2: Boletus eximius Pk., J Mycol 3: Ceriomyces eximius (Pk.) Murrill, Mycologia 1: Tylopilus eximius (Pk.) Singer, Amer Midl Nat 37: Pileus cm broad, at first convex to plano-convex, with age plano-convex, dry to moist; disk even, at first violet brown (11E4), then chocolate brown (6F4); margin even, concolorous with disk; surface matted subtomentose to subvelutinous, becoming subtomentose to tomentose; sterile appendiculate flaps absent. Flesh mm thick, white with a grayish lilac tint to white with pale violet brown tint or pale lilac brown, staining absent; odor mild; taste mild. Hymenophore tubulose, adnexed to depressed. FIGS L. eximium (Halling 7798). 15. Basidiospores. 16. Hymenial cystidia. 17. Caulocystidia. Tubes 5 10 mm long, rarely grayish yellow (4B3) to light brown or brownish purple, becoming grayish brown to dark brown (9F8) to purplish brown (11F8), unchanging when injured; pores 3 per mm, dark brown or purplish brown, then dark pinkish brown, brown when bruised. Stipe (4 )6 15 cm long, 6 12( 20) mm wide, equal to subequal, strict or curved, not radicating, dry; upper half when young finely scabrous, very pale violet, with age subpruinose to scabrous, very pale violet; lower half when young finely scabrous to scabrous, very pale violet, with age scabrous to finely scabrous or subscabrous ridged, very pale violet to lilac or dull lilac gray (16D3); scabers brown; base white, staining not present. Stipe interior solid; flesh above when young pale violet brown to lilac brown, with age pale lilac brown to grayish lilac, staining not present; flesh at base grayish lilac to pale violet brown to pale lilac brown, staining not present. Basal mycelium white. Basidiospores vinaceous brown to pinkish brown, m m, Q 2.55, smooth, ellipsoid to subfusoid to fusoid, inamyloid, with KOH pale brownish yellow. Basidia m m, clavate, hyaline, 4-sterigmate. Hymenial cystidia m 6 8 m, rare, thin-walled, hyaline to goldencolored contents, narrowly fusoid. Tube trama boletoid, hyaline, lateral strata elements m wide, with age subgelatinous. Pileipellis hyphae a trichodermium, in KOH yellow ochraceous, inamyloid; elements m wide, elongated to cylindric or obtuse, encrusted with pigment (but dissolving in KOH), thin-walled, not gelatinized. Pileus trama interwoven, hyaline, inamyloid, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, m 5 14 m wide, cy-

6 HALLING AND MUELLER: COSTA RICAN LECCINUM 493 FIGS L. monticola (Halling 8208). 18. Pileipellis elements. 19. Basidiospores. 20. Caulocystidia. 21. Hymenial cystidia. lindric to clavate to subfusoid, hyaline to brown contents, with encrusting pigment present (with purple acerose crystals dissolving in KOH). Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: solitary to gregarious on soil under Quercus copeyensis, Q. seemannii, Q. rapurahuensis in Costa Rica. Material examined: Costa Rica. Alajuela: Grecia, Grecia. Bosque del Niño, N, W; 1900 m, 29 Jun 1995, Halling 7490 (NY, USJ); 31 May 1996, Halling 7598 (NY, USJ); 15 Jun 1996, Halling 7688 (NY, USJ). San José: Dota, La Chonta, S of Interamerican Highway near km 54 toward Laguna/ Cerro Chonta, N, W, 2400 m, 11 Jul 2000, Halling 8020 (NY, USJ). San Gerardo, Hotel de la Montaña, Savegre, 5 km SW of Cerro de la Muerte, N, W, 2500 m, 20 Jun 1994, Halling 7308 (NY, USJ); 8 Jun 1996, Halling 7643 (NY, USJ). San José: Dota, between km 66/67 of Interamerican Highway, 2.3 km W of highway at Finca El Jaular, N, W; 2234 m, 1 Jul 1998, Halling 7798 (NY, USJ); 5 Jul 2001, Halling 8252 (NY, USJ). The pale violet-to-lilac stipe with fine scabers, brown pileus with purple overtones and brown hymenophore are distinctive local field characters. It is the only species placed in section Eximia by Singer (1986). Originally described from the northeastern U.S.A. (as Boletus robustus Frost non Fr.), L. eximium has been found in the Cordillera Central (Volcan Poas) and the Cordillera Talamanca. The Costa Rican material has a smaller stature and shorter spores (10 13 m) than the forms from the northeastern United States (11 20 m); otherwise they are morphologically identical. Leccinum monticola Halling and G. M. Mueller, sp. nov. FIGS. 4, Etymology. Referring to the montane habitat of the species. Pileus aurantiacus vel brunneo-aurantiacus, fibrillosus vel tomentosus, margine appendiculato et sterili, pileipelle trichodermalis. Contextus albus mutabilis. Stipes albus scabrosus. Basidiosporae in cumulo olivaceo-brunneae subfusiformae parietibus laevibus inamyloideis. Sub Comarostaphylis in montibus Cordillera Talamanca Costa Ricae. HOLOTY PUS (hic designatus): Halling 8288 (USJ). Pileus cm broad, at first obtusely convex, with age plano-convex, dry to moist to subviscid (when wet), even, becoming areolate; disk at first brownish orange (7A-B-C-D8-7-6), then agate brown (6-7E8) to brown to brownish orange to light brown; margin even, concolorous with disk; surface appressed fibrillose to appressed fibrillose scaly to appressed tomentose to tomentose, becoming finely fibrillose; sterile appendiculate flaps present (conspicuous when young); with NH 4 pale brown. Flesh mm thick, white, staining fuscous; odor mild; taste mild. Hymenophore tubulose, free to adnexed. Tubes mm long, pale yellow, becoming light brown to pinkish gray, near bronze-yellow when injured; pores 2 3 mm wide, white, then light brown to pinkish gray, brown when bruised. Stipe (6.5 )10 13( 15) cm long, mm wide, subclavate to clavate, often curved or strict, not radicating, dry; upper half when young scabrous, white, with age scabrous, pale gray, sometimes with pale yellowish-green tints or white; lower half when young scabrous, white, with age scabrous to subglabrous, pale gray, sometimes with pale yellowish-green tints or white; scabers on upper half when young white, with age black to brown, on lower half when young white, with age black to brown; base white, staining not present or blue-green (occasional). Stipe interior solid; flesh above white, staining fuscous; flesh at base white, staining fuscous and/or blue-green. Basal mycelium white. Basidiospores umber brown to olive brown, m m, Q 3.24, smooth, subfusoid to fusoid, inamyloid. Basidia m m,

7 494 MYCOLOGIA FIGS L. neotropicalis (Halling 7216). 22. Pileipellis elements. 23. Basidiospores. 24. Hymenial cystidia. 25. Caulocystidia. Courtesy Kew Bulletin. clavate, hyaline, 4-sterigmate. Hymenial cystidia m 8 11 m, more common toward edge of tubes, thin-walled, hyaline to gray brown contents or honeycolored contents, lageniform to fusoid to ventricose to ventricose rostrate or mucronate, gelatinization absent, encrusting pigment absent. Tube trama boletoid, hyaline, lateral strata elements m wide, subgelatinous (with age). Pileipellis hyphae a trichodermium (often collapsed with age), in KOH brownish, inamyloid; elements m wide, subelongate to elongated, smooth, thin-walled, granular content present, not gelatinized. Pileus trama interwoven, hyaline, inamyloid, with elements m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, m 7 18 m, clavate to subclavate to mucronate or subfusoid (rarely), hyaline or golden yellow contents, with encrusting pigment absent. Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: solitary to scattered or gregarious near Comarostaphylis arbutoides, solely in the páramo, or just below timberline mixed with Q. costaricensis. At present, known only from higher elevation on Volcan Irazu and the Cordillera Talamanca. Material examined: Costa Rica. Cartago: Parque Prusia, W slope of Volcan Irazu, N, W, 2960 m, 20 Jul 1993, Halling 7034 (NY, USJ); 3 Nov 1993, Halling 7122 (NY, USJ); 14 Nov 1993, Halling 7162 (NY, USJ). San José: Dota, San Gerardo, 500 m S of Interamerican Highway on road to San Gerardo, N, W, 3000 m, 21 Oct 1994, Halling 7411 (NY, USJ); 15 Jul 1999, Halling 7886 (INB, NY); 28 Jun 2001, Halling 8208 (NY, USJ); 4 Jul 2001, Halling 8239 (NY, USJ); summit of Cerro de la Muerte, 9 40 N, W, 3491 m, 17 Nov 2001, Halling 8288 (Holotype: USJ; Isotype: NY). Taxa of Leccinum subsection Leccinum are characterized by a conspicuous and distinctive flap of sterile tissue at the margin of the pileus. Furthermore, they frequently have a pileus that is colored a shade of orange or orange brown to red brown and context that changes to pink and/or fuscous when exposed. Many names have been used for species in this subsection, and the taxonomy is in apparent disarray for taxa outside Europe. Mycorrhizal associates in the Northern Hemisphere typically are members of the Betulaceae, or less often of Pinaceae and/or Fagaceae. However, a few species are known to form mycorrhizal associations with members of the Ericaceae (e.g., L. manzanitae Thiers with Arctostaphylos spp. and L. arbuticola Thiers with Arbutus). Leccinum monticola occurs among páramo vegetation on the summit of Cerro de la Muerte and is associated with Comarostaphylis arbutoides. We previously had encountered the fungus commonly at slightly lower elevations ( m) in the Cordillera Talamanca, where Comarostaphylis occurs sympatrically with Quercus costaricensis. It also has been found on Volcan Irazu among similar vegetation. Leccinum neotropicalis Halling, Kew Bull 54: FIGS. 5, Pileus (4 )5 8.5( 13) cm broad, at first convex, with age becoming plano-convex to plane, dry to moist to slippery (when wet), rugose to rugulose to subrugulose; disk even to uneven (rarely), at first brown (6E7) to dark brown (6E-F8) or dark reddish brown (8F8,7,6,5), then remaining brown to dark brown or dark reddish brown; margin subrugulose (rarely), concolorous with disk; surface matted subtomentose to subtomentose to subvelutinous to velutinous; sterile appendiculate flaps absent. Flesh mm thick, white to pale yellow to pale pink, staining absent; odor mild; taste mild. Hymenophore tubulose, adnexed to depressed to deeply depressed. Tubes 7 10 mm long, yellow (3,4A6,5-3B6), becoming greenish yellow (2C6) to olive yellow, unchanging when injured; pores up to 1 mm wide, yellow, unchanging when injured or light cinnamon brown when bruised. Stipe 5.5 9( 11) cm long, mm

8 HALLING AND MUELLER: COSTA RICAN LECCINUM 495 wide, equal to subclavate to clavate, sometimes with a tapered and pointed base, dry; upper half pruinose to ridged with pallid ridges, yellow; lower half when young finely scabrous to ridged with pallid ridges, sordid yellow, with age rarely pink or sordid yellow; scabers on upper half when young gray brown, with age gray brown to tan to clay, on lower half gray brown to tan to clay; base white to pale yellow, staining not present. Stipe interior solid; flesh above white to yellow, staining not present; flesh at base white to yellow, staining not present. Basal mycelium white to yellow. Basidiospores dull cinnamon brown, m m, Q 2.61, smooth, elongate ellipsoid to fusoid to subfusoid, inamyloid. Basidia m 8 12 m, clavate, hyaline, 4-sterigmate. Hymenial cystidia m 6 8 m, thin-walled, hyaline, narrowly lageniform to fusoid to subcylindrical, encrusting pigment absent. Tube trama boletoid, hyaline, in KOH yellowish, lateral strata elements 4 8 m wide, not gelatinized. Pileipellis hyphae hymeniform, pigment encrustations dextrinoid (at pellis/ trama interface); elements m wide, cellular to subisodiametric or subelongate (rarely), thinwalled, often beneath a gelatinized layer. Pileus trama interwoven, hyaline, inamyloid, with elements up to 20 m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, up to 60 m 15 m, clavate to subclavate to subfusoid, occasionally with golden yellow contents, with encrusting pigment absent. Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: Solitary or gregarious on soil under Quercus copeyensis, Q. costaricensis, Q. oocarpa Liebm., Q. rapurahuensis and Q. seemannii in Costa Rica. Material examined: COSTA RICA. Cartago: Guarco, Estrella, 5 km E. of km 31 of Interamerican Highway, N, W, 1685 m, 31 May 1994, Halling 7216 (NY, USJ); 13 Jun 1994, Halling 7281 (NY, USJ). Palo Verde, 4.5kmEofkm31of Interamerican Highway, N, W, 1600 m, 29 May 1996, Halling 7583 (NY, USJ). Tapanti. Parque Nacional Tapanti, Macizo de la Muerte, Area de Conservacion La Amistad Pacifico, N, W, 2600 m, 29 Jun 2001, Halling 8218 (NY, USJ). San José: Dota, Jardín, 3.5 km W of Interamerican Highway at El Empalme, N, W, 2220 m, 26 May 1996, Halling 7554 (NY, USJ). San Gerardo, Hotel de la Montaña, Savegre, 5 km SW of Cerro de la Muerte, N, W, 2500 m, 10 Jun 1994, Halling 7268 (NY, USJ); 27 Jun 2001, Halling 8202 (NY, USJ). La Chonta, S of Interamerican Highway near Cerro Chonta, FIGS L. rugosiceps (Halling 8163). 26. Pileipellis elements. 27. Basidiospores. 28. Caulocystidia. 29. Hymenial cystidia N, W, 2400 m, 11 Jul 2000, Halling 8021 (NY, USJ); 11 Jun 2001, Halling 8170 (NY, USJ). This Leccinum is reminiscent of L. crocipodium (Let.) Watling as currently outlined by Lannoy and Estades (1995) and also would belong in section Luteoscabra. Recent accounts in North America (Smith, Thiers, and Watling 1967; Smith and Thiers 1971) indicate that it occurs in the United States. However, as circumscribed by those authors, that taxon has a much lighter-colored pileus, and perhaps more significant, the context changes with exposure to a pinkish or salmon-to-reddish vinaceous and finally blackish. The Costa Rican material is consistently dark brown to dark red brown with immutable flesh. Leccinum rugosiceps (Pk.) Singer, Mycologia 37: FIGS. 6, Boletus rugosiceps Pk., Bull New York St Mus 94: Krombholzia rugosiceps (Pk.) Singer, Ann Mycol 40: Pileus cm broad, at first convex, with age plano-convex, dry, irregularly pitted to rugulose to wrinkled; disk even, dark brown (7F8) to mustard brown (5E7,6,5) to somalis brown (7E5) to yellowish brown; margin rugulose, concolorous with disk; surface subvelutinous; sterile appendiculate flaps absent;

9 496 MYCOLOGIA with NH 4 no reaction. Flesh mm thick, white, staining pink to pale red or pink followed by fuscous; odor mild; taste mild. Hymenophore tubulose, depressed. Tubes mm long, sordid golden yellow (4C-D5) to wax yellow (3B5); pores up to 1 mm wide, yellow (3A4), then sordid khaki yellow (4D5), brown when bruised. Stipe 7 13 cm long, 7 15 mm wide, equal to subequal to subclavate to clavate, dry; upper half when young pruinose, pale yellow, with age scabrous and ridged with pallid ridges, dark sordid yellow; lower half subscabrous ridged, white to pallid, with age sometimes grayish yellow; scabers on upper half cinnamon to pale yellow, on lower half dark gray to gray; base white, staining not present. Stipe interior solid; flesh white, staining pink; flesh at base white, staining pink. Basal mycelium yellow. Basidiospores m m, Q 3.07, smooth, fusoid to subfusoid, mostly inamyloid (a few dextrinoid), with KOH light brown melleous. Basidia m 9 12 m, clavate, hyaline, 4- sterigmate. Hymenial cystidia m 7 9 m, more common toward edge of tubes, thin-walled, hyaline, subfusoid to fusoid to ventricose, encrusting pigment absent. Tube trama boletoid, hyaline to golden yellow, lateral strata elements m wide, subgelatinous. Pileipellis hyphae hymeniform and forming a palisade (several cells deep), in KOH yellow ochraceous, inamyloid; elements m wide, subelongate to subisodiametric or cellular, smooth, thin-walled, not gelatinized. Intercalary cells isodiametric to subisodiametric, m 7 14 m. Pileus trama interwoven, hyaline, inamyloid, with elements m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, clavate to ventricose to lageniform to fusoid, up to m with pale yellow brown contents or hyaline, with encrusting pigment absent. Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: solitary to gregarious on soil under Quercus copeyensis, Q. rapurahuensis, Q. seemannii (in Costa Rica), Q. humboldtii (in Colombia). Material examined: COSTA RICA. Alajuela: Grecia, Grecia, Bosque del Niño, N, W, 1900 m, 26 Jun 1994, Halling 7344 (NY, USJ); 31 May 1996, Halling 7596 (NY, USJ). Puntarenas: Coto Brus, Las Mellizas, Zona Protectora Las Tablas. Finca La Cafrosa, camino a portones por El Tajo, N, W, 1475 m, 7 Jun 2001, Halling 8139 (NY, USJ). San José: Dota, La Chonta, S of Interamerican Highway near Cerro Chonta, N, W; 2400 m, 16 Jun 1995, Halling 7426 (NY, USJ); 11 Jun 2001, Halling 8172 (NY, USJ). San Gerardo. Albergue de Montaña, Savegre, 5 kmswof FIGS L. tablense (holotype). 30. Pileipellis elements. 31. Basidiospores. 32. Hymenial cystidia. 33. Caulocystidia. Cerro de la Muerte, N, W, 2500 m, 6 Nov 1993, Halling 7131 (NY, USJ); 7 Nov 1993, Halling 7139 (NY, USJ); 10 Nov 1993, Halling 7155 (NY, USJ); 9 Jun 1994, Halling 7258 (NY, USJ); 7 Jun 1996, Halling 7637 (NY, USJ); 8 Jun 1997, Halling 7707 (NY, USJ); 3 Jun 2000, Halling 7978 (NY, USJ); 10 Jun 2001, Halling 8163 (NY, USJ). As the epithet indicates, the pileus surface is rugose, more so in some collections than in others. The hymenophore is pale yellow, and the scabers on the stipe are pale caramel-colored. The context of the basidiomata is white but soon changes to pink or pale red when exposed. It also belongs in section Luteoscabra. Originally described from the eastern United States, L. rugosiceps occurs under Quercus as far south as southern Colombia. In Costa Rica, it seems to be uncommon but widespread in the montane oak forests in the Cordillera Talamanca and Cordillera Central (Volcan Poas). Leccinum tablense Halling and G. M. Mueller, sp. nov. FIGS. 7, Etymology. Referring to the locality where the species was first discovered. Pileus griseo-brunneus interdum maculis albis aequus vel foveatus irregulariter vel areolatus, margine non-appendiculato, hyphis pileipellis cellulosis et hymeniformis, trama pileo et stipitis immutabilis, caulocystdiis orientibus hyphis

10 HALLING AND MUELLER: COSTA RICAN LECCINUM 497 isodiametricis. Sub Quercus in montibus Cordillera Talamanca Costa Ricae. HOLOTY PUS (hic designatus): Halling 8136 (USJ). Pileus cm broad, at first convex, with age plano-convex, moist, even or irregularly pitted at first, remaining irregularly pitted and sometimes becoming areolate; light grayish brown or tan (6E6-5-4), sometimes with irregularly distributed white zones; margin even; surface subvelutinous, becoming subtomentose; sterile appendiculate flaps absent. Flesh 6 8 mm thick, white, staining absent; odor mild; taste mild. Hymenophore tubulose, depressed. Tubes 9 14 mm long, white, becoming orangish white (5A2), unchanging when injured; pores 1 mm wide, white, then orangish white, unchanging when injured. Stipe 5 8 cm long, 5 8 mm wide, equal to subclavate, strict, dry; upper half when young finely scabrous, white, with age scabrous, white; lower half when young subscabrous ridged, white, with age subscabrous-ridged to scabrous, white; scabers on upper half when young gray to black, with age black, on lower half when young dark gray or black, with age black; base white, staining not present. Stipe interior solid or soon hollow; flesh above white, staining not present, with age whitish yellow at base, staining not present. Basal mycelium white. Basidiospores m, Q 2.83, smooth, fusoid to subfusoid, mostly inamyloid (a few dextrinoid), light brown melleous in KOH. Basidia m, clavate, hyaline in KOH, lightly dextrinoid, 4-sterigmate. Hymenial cystidia m, more common toward edge of tubes, thinwalled, hyaline, subfusoid to narrowly fusoid, encrusting pigment absent. Tube trama boletoid, hyaline to golden yellow, lateral strata elements m wide, subgelatinous. Pileipellis hyphae hymeniform and forming a palisade (several cells deep), in KOH yellow ochraceous, inamyloid; elements m wide, subelongate to subisodiametric or cellular, smooth, thin-walled, not gelatinized. Intercalary cells isodiametric to subisodiametric, m broad. Pileus trama interwoven, hyaline, inamyloid, with elements m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, clavate to subclavate, or irregularly ventricose, typically subtended by isodiametric cells, with pale gray brown contents or hyaline, with encrusting pigment absent. Stipe trama hyphae parallel, cylindric, hyaline, inamyloid. Clamp connections absent. Habit, habitat and distribution: gregarious on soil under Quercus seemannii in the Cordillera Talamanca. Material examined: Costa Rica. Puntarenas: Coto FIGS L. talamancae (Halling 7142). 34. Pileipellis elements. 35. Basidiospores. 36. Hymenial cystidia. 37. Caulocystidia. Courtesy Kew Bulletin. Brus, Las Mellizas, Zona Protectora Las Tablas, Finca La Cafrosa, camino a portones por El Tajo, N, W; 1475 m, 7 Jun 2001, Halling 8136 (Holotype: USJ; Isotype: NY). Leccinum tablense vaguely recalls features of L. talamancae, but unlike that taxon it does not exhibit any oxidation reactions in the exposed flesh: pink to reddish orange in the pileus and upper part of the stipe and the blue-green to deep blue in the lower part of the stipe. The hymeniform nature of the pileipellis also is reminiscent of L. talamancae, but in lieu of a description of fresh material they can be distinguished by spore size ( m for L. tablense, and m for L. talamancae). Leccinum tablense also would fit the circumscription of subsection Pseudoscabra as recently outlined by Lannoy and Estades (1995). Leccinum talamancae Halling, Gómez, and Lannoy in Halling, Kew Bull 54: FIGS. 8, Pileus 2 6( 8) cm broad, at first convex, with age plano-convex, dry to moist to viscid (in wet weather), even, becoming areolate to pitted to rugose to rugulose to wrinkled; disk even or rugulose or wrinkled, at first brownish gray to vandyke brown, then cocoa brown to vandyke brown; margin even or rugulose or wrinkled, when young cocoa brown to dark brown, then cocoa brown to dark reddish brown; surface subtomentose to matted subtomentose, becoming matted subtomentose to matted tomentose; sterile appendiculate flaps absent; with KOH orange brown to yellow. Flesh up to 8 mm thick, white, staining slowly pink to reddish orange; odor mild; taste mild. Hymenophore tubulose, adnexed to depressed to

11 498 MYCOLOGIA deeply depressed. Tubes 8 15 mm long, yellowish white, becoming brownish orange to mustard yellow, avellaneous when injured; pores up to 1 mm wide, white, dark reddish brown when bruised. Stipe 4 14 cm long, (4 )10 20 mm wide, equal to subclavate to clavate, strict or curved, dry; upper half when young finely scabrous and finely subpruinose ridged, white, with age finely scabrous to subpruinose, white; lower half when young scabrous, white, with age subscabrous ridged, white; scaber on upper half when young gray, with age black, on lower half when young gray, with age black; base white, staining blue-green to deep blue. Stipe interior solid; flesh above when young white, with age white, staining pink to pinkish brown or reddish orange; flesh at base when young white, with age white, staining blue or greenish blue to dark blue; with NH 4 bright grass-green at stipe base. Basal mycelium white, changing to blue when picked. Basidiospores m, mean Q 3.5, smooth, elongate ellipsoid to subfusoid, inamyloid. Basidia m, clavate, hyaline, 4- sterigmate. Hymenial cystidia m, moderately abundant, thin-walled, sometimes with gray brown contents or hyaline, lageniform to clavate-ventricose to subfusoid to clavate. Tube trama boletoid, hyaline, lateral strata elements 4 8 m wide, subgelatinous. Pileipellis hyphae hymeniform (several cells in depth), inamyloid; elements up to 75 m wide, cellular or subisodiametric and in sphaerocyst-like chains or subelongate, smooth or encrusted with pigment, thin-walled. Intercalary cells isodiametric to subisodiametric or subcylindric (rarely). Pileus trama interwoven, hyaline, inamyloid, with elements up to 20 m wide, smooth, thin-walled. Stipitipellis hyphae vertically oriented, parallel, giving rise to clusters of caulocystidia, m, subfusoid to clavate to subclavate or lageniform (rarely), often with brown contents. Clamp connections absent. Habit, habitat and distribution: solitary to gregarious on soil under Quercus copeyensis, Q. costaricensis, Q. seemannii (in Costa Rica), Q. humboldtii (in Colombia, fide Franco-Molano et al 2000). Material examined: Costa Rica. Cartago: El Guarco, Tapanti, Parque Nacional Tapanti, Macizo de la Muerte, Area de Conservacion La Amistad Pacifico, N, W, 2600 m, 29 Jun 2001, Halling 8221 (NY, USJ). San José: Dota, La Chonta, S of Interamerican Highway near Cerro Chonta, N, W, 2400 m, 7 Jun 1994, Halling 7239 (NY, USJ); 19 Jun 1994, Halling 7296 (NY, USJ); 6 Jun 1996, Halling 7627 (NY, USJ); 11 Jun 2001, Halling 8168 (NY, USJ). San José: Perez Zeledón, Villa Mills, C.A.T.I.E. Experimental Forest, N, W, 2880 m, 8 Nov 1993, Halling 7148 (NY, USJ); 25 Nov 1993, Halling 7197 (NY, USJ); 22 Jun 1994, Halling 7335 (NY, USJ); 10 Jun 1997, Halling 7719 (NY, USJ). San José: Dota, Jardín, 3.5 km E of Interamerican Highway at Empalme, N, W, 2220 m, 5 Jun 1996, Halling 7617 (NY, USJ). San Gerardo, Albergue de Montaña, Savegre, 5 kmswofcerro de la Muerte, N, W, 2500 m, 10 Jun 1994, Halling 7269 (NY, USJ); 19 Oct 1994, Halling 7392 (NY, USJ); 8 Jun 1996, Halling 7647, 7648 (NY, USJ); 10 Jul 2000, Halling 8001 (NY, USJ); 1.5 km from Interamerican Highway on road toward San Gerardo, N, W, 2860 m, 24 Jun 1995, Halling 7475 (NY, USJ). The flesh is white at first when exposed, then becomes pink to reddish orange in the pileus and stipe apex. In mid-stipe the change is to a greenish blue and a deep, dark blue in the base of the stipe. These oxidation reactions appear to be consistent with this taxon. The pileus color seems to vary from a caramel to dark gray-brown and typically is rugulose to rugose/wrinkled, depending on individual collections. KOH is yellow to orange brown on the pileus surface. In a recent monograph of European Leccinums (Lannoy and Estades 1995), the Costa Rican material comes closest to L. variicolor Watling in subsection Pseudoscabra. However, that species is apparently restricted to an association with Betula and the pileipellis elements are cylindrical in shape not isodiametric. At present, L. variicolor is unknown in the Americas, even though mycorrhizal host shifts have been documented and described for other agarics (e.g., Mueller and Strack 1992, Bougher and Malajczuk 1985, Bougher et al 1994). Clearly, more collecting in temperate and tropical America will provide a better picture of species composition and mycorrhizal associates. LITERATURE CITED Bidartondo MI, Bruns TD Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure. Molec Ecol 10: Bougher NL, Malajczuk N A new species of Descolea (Agaricales) from western Australia, and aspects of its mycorrhizal status. Austral J Bot 33: , Fuhrer BA, Horak E Taxonomy and biogeography of Australian Rozites species mycorrhizal with Nothofagus and Myrtaceae. 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