Elaphomyces (Ascomycota, Eurotiales, Elaphomycetaceae)
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New Zealand Journal of Botany Vol. 50, No. 4, December 2012, 423Á433 Sequestrate fungi of New Zealand: Elaphomyces (Ascomycota, Eurotiales, Elaphomycetaceae) Michael A Castellanoa*, Ross E Beeverb$ and James M Trappec aUS Department of Agriculture, Forest Service, Corvallis, OR, USA; bManaaki Whenua Landcare Research, Auckland, New Zealand; cDepartment of Forest Ecosystems and Society, Oregon State University, Corvallis, OR, USA (Received 6 January 2012; accepted 16 August 2012) Four species of the sequestrate fungal genus Elaphomyces are reported from New Zealand: Elaphomyces bollardii sp. nov. associated with Leptospermum spp. and Kunzea ericoides, E. luteicrustus sp. nov. associated with Nothofagus menziesii, E. putridus sp. nov. associated with Nothofagus spp., and an unnamed species associated with Nothofagus spp. Keywords: biodiversity; systematics; Eurotiales; Elaphomycetaceae; Elaphomyces bollardii; Elaphomyces luteicrustus; Elaphomyces putridus; New Zealand Introduction we currently accept c. 55 species as valid and Sequestrate fungi comprise those macrofungal distinct. The genus is widespread across the taxa in which the spores mature in a more or northern hemisphere including Europe, Asia less enclosed sporocarp, with spores that are (Japan, China & Singapore) and North America, usually not forcibly discharged, and in which and the southern hemisphere including South the sporocarp itself is usually indehiscent. Most America (Argentina and Guyana), Central taxa are hypogeous, producing their sporocarps America (Costa Rica and Mexico), Australia, beneath the soil surface, although some are New Zealand and Papua New Guinea. Castellano emergent or epigeous. Sequestrate fungi occur et al. (2011) recently described 13 new within the Agaricomycotina, Glomeromycotina, Elaphomyces species from Australia including Mucoromycotina and Pezizomycotina (includ- reassignment of a few collections that had been ing the truffles sensu stricto) (Castellano & attributed to Elaphomyces species names from Trappe 1990, 1992; Blackwell et al. 2006). As Europe (Cooke 1892; Rodway 1918; Dodge part of ongoing studies of sequestrate Pezizo- 1929). Recent field-work focusing on sequestrate mycotina in New Zealand (Trappe 1979; fungi reveals Elaphomyces is commonly encoun- Trappe et al. 1992) we have examined the genus tered in New Zealand and Australia and has also Elaphomyces, in the Elaphomycetaceae. been collected in Papua New Guinea (Castellano Index Fungorum (2011) lists 98 Elaphomyces and Trappe, unpublished data). The New names of taxa ectomycorrhizal with a variety Zealand material comprises three new species of vascular plant hosts. Castellano is currently of Elaphomyces and a putative fourth as yet re-evaluating all species of Elaphomyces and undescribed Elaphomyces species. *Corresponding author. Email: [email protected] $Ross E Beever (1946Á2010). ISSN 0028-825X print/ISSN 1175-8643 online # 2012 The Royal Society of New Zealand http://dx.doi.org/10.1080/0028825X.2012.725057 http://www.tandfonline.com 424 MA Castellano et al. Molecular evidence places Elaphomyces Elaphomyces Nees:Fr. Syst. Mycol. 3: 57 (1829) in family Elaphomycetaceae, order Eurotiales (Lumbsch & Huhndorf 2007). Elaphomyceta- TYPE SPECIES. Elaphomyces granulatus Fr. Syst. ceae as presently conceived contains two gen- Mycol. 3: 58 (1829) era, Elaphomyces and Pseudotulostoma (Miller et al. 2001). Recent molecular analysis evidence Ascomata globose to subglobose or irregular, indicates that Pseudotulostoma is nested with (5 )10Á30 (40) mm broad, surface brown to Elaphomyces and should probably be considered dark blue or black, smooth or variously warty asynonymofElaphomyces (Reynolds 2001). or bumpy, often enclosed in a crust of mycelia, ectomycorrhizae, debris and soil particles. Peri- Methods dium (0.5 )2Á4(5) mm thick, fleshy to Macroscopic descriptions refer to fresh material leathery, sometimes carbonaceous; usually hard unless specified. Colour designations follow com- and brittle when dried except in one species. mon usage or the ISCC-NBS Centroid Color Gleba a single chamber, often hollow in youth Charts (Kelly & Judd 1965) for which the colour but usually becoming filled with ascogenous block number is given in parentheses. Specimens and associated cottony hyphae, and finally with were examined in 3% KOH, Melzer’s reagent and a more or less powdery slate blue, purpleÁ cotton blue. Microscopic descriptions are based brown, dark brown, greyÁblack or black spore on 3% KOH mounts unless specified. Spore mass, nearly hollow with glebal remnants measurements are based on 20 spores from the adhering to the inner peridial surface in one holotype collection. Spore dimensions include species. Asci globose to subglobose, thin-walled ornamentation. Dried spores were mounted on or thick-walled, evanescent before spore ma- pegs with double-sided tape and coated with gold turity, non-amyloid, with 1Á8 spores. Ascos- for scanning electron microscopy (SEM) exam- pores subglobose to globose, 8Á65 mm broad ination with an AmRay 3300 FE field emission including ornamentation, often hyaline in scanning electron microscope. Herbaria are ab- youth, at maturity olive, redÁbrown to dark breviated according to Index Herbariorum (2011). brown or nearly black, ornamentation various, Unless indicated, descriptions refer to usually of spines or rods aggregated in various mature sporocarps with fully developed gleba. shapes (clumps, ridges, lines), sometimes Sporocarps in which the inner layer of the slightly to coarsely verrucose or with a partial peridium has collapsed against the carbonac- to full reticulum in surface view, non-amyloid eous outer layer are referred to as over-mature. and non-dextrinoid. Taxonomy Key to New Zealand species of Elaphomyces 1. Peridium splitting open (like a Geastrum), epicutis with yellow tones and leathery, spores (22 )24Á28 (29) mm broad, associated with Nothofagus ....................Elaphomyces putridus Peridium remaining closed, epicutis black and carbonaceous, spores 525 mm broad, associated with Leptospermum or Nothofagus ...........................................................................................2 2. Spores 22Á25 mm broad, coarsely reticulate, associated with Nothofagus ........Elaphomyces sp. ...................................................................................................................... (‘novaeÁzelandiae’) Spores 512.5 mm broad, verrucose, associated with Leptospermum or Nothofagus ...............3 3. Crust mycelium orange, epicutis of polygonal warts when dry, spores (9 )10Á12 mm broad, associated with Leptospermum ................................................................. Elaphomyces bollardii Crust mycelium brilliant yellow, epicutis smooth when dry, spores (10 )11Á12.5 mm broad, associated with Nothofagus..... .............................................................Elaphomyces luteicrustus Sequestrate fungi of New Zealand 425 Figure 1 Elaphomyces bollardii, Beever 175. A, Ascomata surface with adherent mycelium removed. B, Peridial cross-section, ol outer peridial layer, nl inner peridial layer, g gleba. C, Asci with 5 spores visible. D, Ascospores, photomicrograph cross-section showing edge detail. E, Electron micrograph of ascospores. F, Electron micrograph showing ornamentation detail on ascospores. Scales A 300 mm; B 1 mm; C 30 mm; D 12 mm; E 6 mm; F 6 mm. ETYMOLOGY. From Greek elaph (deer) and myces hyphae, very dark brown to mostly black; fungus, in reference to the observation of deer enclosed in a crust up to c. 10 mm thick of scraping the soil for Elaphomyces sporocarps. conspicuous vivid orange (#48) to vivid red (#34) hyphae intermingled with soil, debris and Elaphomyces bollardii Beever, Castellano & occasional roots, the hyphae drying greyÁredÁ Trappe, sp. nov. Fig. 1AÁ1F orange (#39) to strong redÁorange (#35). Peridium 2-layered, 1.45Á2.9 mm thick: peridial MB 801200 outer layer 150Á200 mm thick, carbonaceous, black or occasionally with dark brown patches; Ascomata globose to subglobose, occasionally peridial inner layer 1.3Á2.7 mm thick, uniform, lobed, 7Á22 mm broad. Peridial surface smooth white with occasional traces of strong red (#12) and mottled with areas of tightly appressed to strong purpleÁred (#255) abutting the gleba 426 MA Castellano et al. and sporadically elsewhere especially when coral-like, labyrinthine ridges. Spore colour in immature, soft with the texture of Styrofoam, KOH and Melzer’s reagent greenÁgrey in youth, reduced to a thin, yellowÁwhite (#92), parch- golden brown when mature. ment-like layer when over-mature. Gleba pow- dery, dark grey (#266) to black (#267), with HOLOTYPE. AUCKLAND: Waitakere Ranges, distinct off-white dissepiments arising from the Huia, Parau track, R.E. Beever 175,28Oct1982 inner layer and ramifying up to 0.5 mm into the (PDD 43292, isotype OSC). powdery spore mass. Odour nil. Taste slightly nutty tending to peppery. HABIT, HABITAT AND SEASON. Hypogeous up to Crust hyphae 1.5Á3(4) mm diam, thin- 10 cm or more deep in mineral soil, rarely walled, loosely packed, sparingly branched, partially exposed, over-mature and perhaps sparingly septate, pinkÁbrown in KOH, redÁ years-old specimens sometimes emergent by brown and encrusted with particles in water. erosion, gregarious in shrub-land and second- Peridial outer layer of very dark brown, com- growth podocarpÁbroadleaf forest, putatively pact hyphae, 2.5Á5 mm broad, walls 1Á1.5 mm mycorrhizal with Kunzea ericoides (A. Rich.) thick, forming a textura oblita. Peridial inner