New Zealand Journal of Botany Vol. 50, No. 4, December 2012, 423 433
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 (1946 2010).
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 )10 30 ( 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 )2 4( 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, purple cotton blue. Microscopic descriptions are based brown, dark brown, grey black 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 1 8 spores. Ascos- for scanning electron microscopy (SEM) exam- pores subglobose to globose, 8 65 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, red brown 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 )24 28 ( 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 22 25 mm broad, coarsely reticulate, associated with Nothofagus ...... Elaphomyces sp...... (‘novae zelandiae’) Spores 512.5 mm broad, verrucose, associated with Leptospermum or Nothofagus ...... 3 3. Crust mycelium orange, epicutis of polygonal warts when dry, spores (9 )10 12 mm broad, associated with Leptospermum ...... Elaphomyces bollardii Crust mycelium brilliant yellow, epicutis smooth when dry, spores (10 )11 12.5 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 grey red Trappe, sp. nov. Fig. 1A 1F orange (#39) to strong red orange (#35). Peridium 2-layered, 1.45 2.9 mm thick: peridial MB 801200 outer layer 150 200 mm thick, carbonaceous, black or occasionally with dark brown patches; Ascomata globose to subglobose, occasionally peridial inner layer 1.3 2.7 mm thick, uniform, lobed, 7 22 mm broad. Peridial surface smooth white with occasional traces of strong red (#12) and mottled with areas of tightly appressed to strong purple red (#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 green grey in youth, reduced to a thin, yellow white (#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 1.5 3( 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, pink brown in KOH, red 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 podocarp broadleaf forest, putatively pact hyphae, 2.5 5 mm broad, walls 1 1.5 mm mycorrhizal with Kunzea ericoides (A. Rich.) thick, forming a textura oblita. Peridial inner Joy Thomps. or Leptospermum scoparium J.R. layer composed of sparingly septate, hyaline Forst. & G. Forst. Found August through hyphae, 2 5 mm broad, walls 90.5 mm thick, November and also in January. in densely packed bundles next to the epicutis forming a textura porrecta, near the gleba more DISTRIBUTION. New Zealand, North Island. loosely packed and anastomosing, irregularly inflated to 10 mm broad, and forming a ETYMOLOGY. For plant physiologist E.G. textura intrica, septa not reacting with Melzer’s Bollard, scientific mentor to REB and supporter reagent. Gleba with occasional, sparingly of mycology during his tenure as Director of branched, sparingly septate, thin-walled hy- Plant Diseases Division. phae, 1 3 mm broad, sometimes inflated to 15 mm broad. Asci (4 ) 8-spored, globose, eva- REMARKS. Elaphomyces citrinus Vittad. from nescent, 20 30 mm broad, hyaline, with a short, Europe and E. singaporensis Corner & Hawker hyaline, cone-like or cylindrical stalk 3 5 mm from Singapore differ in their yellow crust long by 4 5 mm broad, thin-walled; arising hyphae and larger spores (12 16 mmbroad). from thin-walled, pale green to hyaline asco- Elaphomyces viridiseptum Trappe & Kimbr. from genous hyphae in knots, 4 5 mm broad. Ascos- the southeastern USA differs by its green septal pores globose, (9 )10 12 mm broad (mean hyphae in Melzer’s reagent and smaller spores 11.1 mm) including ornamentation, with light (8 10 mm broad). See the discussion for E. microscopy ornamentation in spore surface view luteicrustus for similar species from Australia. appearing finely punctate or verruculose; in The two Australian species (E. rugosisporus cross-sectional view spore ornamentation ap- Castellano, Trappe & Vernes and E. symeae pearing two layered, inner layer a palisade of Castellano, Trappe & Lebel) with orange to red pale brown, fine spines, 1 1.5 mm tall, outer layer mycelium enveloping the sporocarp have a continuous, dark brown, thin (B 0.5 mmthick) slightly larger spores and differ in spore orna- tectum; SEM reveals the inner layer as a palisade mentation. The vivid orange or red crust of of crowded, fine spines (B 0.25 mmbroad), hyphae enclosing the ascoma differs from all overlain by a distinct tectum which is mostly known Asian Elaphomyces species. continuous but sometimes with irregularly cir- One site contained c. 500 specimens spread cular to elongate pits 0.2 0.6 mm across, the across an area of 25 m2 on a slope colonized by surface of the tectum appearing as low, fine, orange yellow mycelium. Sequestrate fungi of New Zealand 427
PARATYPES. AUCKLAND: Waitakere Ranges, sparingly branched, loosely packed, straight Huia, Parau track, R.E. Beever 176, 20 Oct 1982 or sinuous; hyaline to pale yellow, smooth. (PDD 43293, OSC); R.E. Beever s.n., 17 Sep Peridial outer layer of dark brown, isodiametric 1977 (PDD 37116, OSC); R.E. Beever, 25 Sep to elongate hyphae, 4 9 mm broad, walls 91 mm 1977 (PDD 37130); R.E. Beever s.n., 19 Aug thick, forming a combination of textura 1978 (PDD 38495) and R.E. Beever & J.M. prismatica and textura angularis. Peridial inner Trappe 6988 (PDD 99963, OSC); Titirangi, M. layer near the outer layer composed of com- Hodgkins, Nov 1931 (PDD 29705, OSC); M. pact, hyaline, mostly isodiametric hyphae, 5 7 Hodgkins, 1931 (PDD 29706, OSC); Little Huia, mm broad, walls 1 2 mm thick, near the gleba P. Turner, Nov 1987 (PDD 100534). Kaipara less densely packed and up to 12 mm broad, Harbour, Kaukapakapa Estuary Scientific Re- forming a combination of textura oblita and serve, R.E. Beever, 22 Sep 1985 (PDD 88003). textura angularis. Gleba with hyaline, sinuous, COROMANDEL: Little Barrier Island, R.E. sparingly septate, thin-walled hyphae, 2 4 mm Beever 37, 23 Jan 1980 (PDD 43287, OSC). broad, occasionally swollen to 6 mm. Asci (7 ) 8-spored, globose, collapsing as spores mature, Elaphomyces luteicrustus Beever, Castellano & 23 27 mm broad, hyaline at first then darker as Trappe, sp. nov. Fig. 2A 2F spores mature, thin-walled, stalk indistinct, arising from knots of ascogenous, hyaline, MB 801201 contorted hyphae up to 4 mm broad, walls 91 mm thick. Ascospores globose, (9 )10 12 Ascomata globose, subglobose to irregular, ( 12.5) mm broad (mean 11.2 mm), including 11 17 mm broad; surface completely enclosed ornamentation, with light microscopy orna- in a persistent husk of tightly adherent, inter- mentation in spore surface view appearing twined ectomycorrhizae which gives rise to a coarsely warty; warts dark brown, subcircular thick (95 mm) layer of roots, brilliant yellow to irregular in surface view, appearing some- (#83) mycelium, soil and debris that forms a whatroundedtoirregularattheapexinside crust surrounding each ascoma. Peridial sur- view, separated by narrow, pale brown, face (after removal of ectomycorrhizal husk) vermicular striae; in cross-sectional view black, with verrucae circular to polygonal or spore ornamentation appearing as coarse, irregular in face view, mostly flat-topped or truncate rods, 1 2 mm tall; SEM reveals these slightly rounded, 100 150 mmbroad,950 mm wartsorrodstobebundlesoffewtomany tall; crust hyphae drying yellow white (#92). individual spines fused together from their base 3 Peridium 2-layered, 1.5 2.5 mm thick: outer to c. 4 their length, spine apices digitate, the layer, thin, 40 100 mm thick, slightly wavy in spines mostly flaring away from each other at section, black, carbonaceous; inner layer up to the tip to form a corona, the bundles irregularly 3.4 mm thick, distinctly demarcated from shaped from nearly circular to elongated ridges; outer layer, white to grey white, soft, uniform in KOH and Melzer’s reagent spores pale olive in colour and texture. Gleba powdery, dark green grey to brown. grey (#156) when fresh, drying grey black (#266), with numerous pale grey dissepiments arising HOLOTYPE. TAUPO, Kaimanawa Forest Park, from inner peridial layer and extending into the Kiko Rd, Ngapuketurua Track, 7 May 1987, powdery spore mass giving the sectioned gleba a R.E. Beever 707 (PDD 56293, isotype OSC). stringy appearance. Odour nil. Taste not noted. Crust hyphae walls 0.5 1.0 mmthick,septate, HABIT, HABITAT AND SEASON: Hypogeous, gre- 3 5 mm broad, sometimes swollen in bulbous garious in duff under Nothofagus menziesii fashion on one side of the septa to 7 mm, (Hook. f.) Oerst.; May. 428 MA Castellano et al.
Figure 2 Elaphomyces luteicrustus, Beever 707. A, Ascomata surface with adherent mycelium removed. B, Peridial cross-section, ch crust hyphae, ol outer peridial layer, nl inner peridial layer, g gleba. C, Asci with 5 spores visible. D, Ascospores, photomicrograph showing surface and edge detail. E, Electron micrograph of ascospores. F, Electron micrograph showing ornamentation detail on ascospores. Scales A 300 mm; B 1 mm; C 25 mm; D, E 11 mm; F 2 mm.
DISTRIBUTION. New Zealand, North Island, of the crust hyphae, the appearance of the spore known only from the type locality. ornamentation in cross-section, and the host asso- ciation. It somewhat resembles E. citrinus and ETYMOLOGY.FromLatinluteus, bright yellow, E. singaporensis from the northern hemisphere and crustus, crust, in reference to the colour of but they differ in their larger spores (12 16 mm the crust mycelium. broad). It also resembles E. viridiseptum but the latter has green septal hyphae in Melzer’s reagent REMARKS. This species resembles E. bollardii and smaller spores (8 10 mm broad). It also also from New Zealand, but differs in the colour resembles the Australian E. aurantius Castellano, Sequestrate fungi of New Zealand 429
Trappe & Vernes, E. cooloolanus Castellano, Crust hyphae not observed. Peridial outer Trappe & Vernes, E. pedicellaris Castellano, layer not easily discernable. Inner layer not Trappe & Vernes, E. rugosisporus and E. symeae observed. Glebal hyphae thin-walled, 2 3 mm either in spore size or colour of the mycelium broad, septate. Asci not observed. Ascospores enclosing the sporocarp. Both E. aurantius and globose, (18.5 )20 24 ( 25) mm broad E. pedicellaris may have yellow encrusting my- (mean 22.0 mm) including ornamentation, celium surrounding the sporocarp but the with light microscopy the spore surface view spores of E. aurantius are larger (14 16 mm appearing as a complete alveolate reticulum, broad, mean 15.0 mm) and the spore orna- the reticular walls dark brown, 91.0 mm thick, mentation coarser, while the distinctive pedicel- the spore surface within the walls pale brown, late spore ornamentation of E. pedicellaris easily 1 4 mm broad; in cross-sectional view the spore separates it from E. luteicrustus. Elaphomyces ornamentation falsely appearing as dark cooloolanus, E. rugosisporus and E. symeae have brown, coarse, truncate warts, 2 3 mm tall; similar-sized spores but they all have differently SEM reveals the ornamentation to be a com- coloured mycelium (orange to red in plete alveolate reticulum as observed with light E. rugosisporus and E. symeae and white to tan microscopy in the spore surface view, the for E. cooloolanus) than the yellow that sur- alveolae 4 5-sided and 2 5 mm broad, the rounds E. luteicrustus sporocarps; the spore alveolar walls 1 4 mm tall and B1.0 mm broad ornamentation differs significantly between the with somewhat digitate margins; spores dark four species. The brilliant yellow crust of hyphae olive brown to black in KOH, golden brown in enclosing the ascoma differs from all known Melzer’s reagent. Asian Elaphomyces species. HABIT, HABITAT AND SEASON. Hypogeous Elaphomyces sp. ‘novae-zelandiae’ Fig. 3A 3F amongst roots and litter in Nothofagus forests, over-mature sporocarps sometimes exposed Only over-mature, disintegrating sporocarps by erosion, putatively mycorrhizal with have so far been found for this species, which Nothofagus menziesii, N. fusca (Hook. f.) Oerst. can be readily distinguished by spore characters and N. solandri (Hook. f.) Oerst.; April, May from the other New Zealand taxa. Because the and October. description lacks many details we do not for- mally name it. The informal name ‘novae- DISTRIBUTION. New Zealand, North Island and zelandiae’ reflects its distribution in both the South Island. North and South Islands. REMARKS. This Elaphomyces species differs Ascomata globose to subglobose, occasionally from the others of New Zealand and Asia in pyriform, 5 18 mm broad; traces of red brown its conspicuously reticulated spores. This fea- to brown mycelium present on peridial surface. ture is shared with only a few other species in Peridial surface black, smooth, sometimes the genus, and they only occur in the northern eroded to appear finely warty, in cross-section hemisphere. Elaphomyces persoonii Vittad. dif- 0.1 0.2 mm thick, 2-layered: Peridial outer layer fers in its coarsely verrucose peridium and black, carbonaceous, up to 0.2 mm broad. conspicuous basal tuft of yellow mycelium; Peridial inner layer brown, adhering in fragments E. cyanosporus Tul. & C. Tul. in its papillate to the outer layer and gleba. Gleba powdery, peridium and basal tuft of white mycelium; and dark brown black (#65), flecked with off-white E. reticulosporus B.C. Zhang in it smaller (17 hyphae, sometimes detached and lying loose in 22 um) spores (Zhang & Minter 1989). No the outer layer of the peridium. Odour not species described before now from the Southern recorded. Taste not recorded. Hemisphere have a distinct, coarse reticulum. 430 MA Castellano et al.
Figure 3 Elaphomyces sp. ‘novae-zelandiae’, Beever 169. A, Electron micrograph showing ornamentation detail on ascospores. B, Electron micrograph of ascospore. C, Ascospores, photomicrograph showing surface detail. D, Ascospores, photomicrograph cross-section showing edge detail. E, Ascomata surface with adherent mycelium removed. Scales A 5 mm; B 10 mm; C, D 22 mm; E 300 mm.
COLLECTIONS EXAMINED. TAUPO: Kaimanawa Elaphomyces putridus Beever, Castellano & Forest Park: Clements Rd., R.E. Beever 169,2 Trappe, sp. nov. Fig. 4A 4D Oct 1982 (PDD 43291); Kiko Rd, Ngapuketur- MB 801202 ua track, R.E. Beever 710, 7 May 1987 (PDD 56295). BULLER: Murchison, Warbeck Scenic Ascomata globose to depressed globose, 12 22 Reserve, Maruia Saddle, R.E. Beever 112,16 mm broad, apex usually slightly indented, and April 1983 (PDD 56294). FIORDLAND: indistinctly radially furrowed, the peridium Fiordland National Park, Waiau River, Rain- breaking at maturity along the furrows and bow Reach, Kepler track, R.E. Beever 1007,18 reflexed to form more or less 5 irregular rays May 1990 (PDD 57838). and expose the spore mass; surface glabrous to Sequestrate fungi of New Zealand 431
Figure 4 Elaphomyces putridus, Trappe 12572. A, Ascomata showing surface and in cross-section. B, Ascomata showing dehiscence with patches of glebal tissue adhering to inner peridial layer. C, Peridial cross- section. D, Ascospores, photomicrograph showing surface detail. E, Electron micrograph of ascospores. F, Electron micrograph showing ornamentation detail on ascospores. Scales A 12 mm; B 18 mm; C 700 mm; D 25 mm; E 12 mm; F 2 mm. minutely scurfy, the scruffiness due to surface coarsely verrucose with irregular warts low (950 mm tall), wide (9100 mm broad), with dark brown apices. Gleba initially hollow, evanescent bumps, pale grey yellow to pale becoming more or less packed with clumped, yellow brown; associated with cream-coloured aggregated to slightly powdery black spore mass. ectomycorrhizae and yellow mycelium but lack- Taste mild. Odour initially faint, smelling of ing a well-developed crust. ETOH, FeSO4,KOH rotting carrion when sporocarp opens. negative on surface and spore mass. Peridium 2- Associated hyphae 3 5 mm diam, sparingly layered, 2 3mmthick(1 1.3 mm thick when branched and septate, thin-walled, pale yellow. dried); peridial outer layer thin, concolorous with Peridial outer layer 90 110 mm, of thin-walled surface; peridial inner layer white, crisp, its inner (90.5 mm), elongate (25 60 7 25 mm) to 432 MA Castellano et al. isodiametric (10 18 mm) cells, hyaline to pale geous habit of the ascomata at maturity, and by yellow near surface, forming a textura angularis to the splitting of the peridium from the apex into textura prismatica. Peridial inner layer 2 3 mm, of reflexed rays similarly to Geastrum spp. to raise thin-walled (90.5 mm), elongate cells, 50 90 8 the ascoma partly out of the ground. Both known 15 mm, septa 3 10 mm broad, forming a textura collections were made during rainy weather, but intrica, except the warts lining the glebal cavity we do not know whether such conditions are which comprise isodiametric cells, 9 18 mm essential for dehiscence. We hypothesize that broad, forming a textura angularis to globosa. sporocarp dehiscence coupled with development Glebal spore mass with occasional hyaline, spar- of a carrion-like odour and aggregated rather ingly branched hyphae, 1.5 2 mmbroad.Asciwith than powdery spore mass at maturity are asso- (4 ) 8 spores, hyaline, globose, 35 50 mmbroad, ciated with dispersal by insects such as flies, but becoming irregular as spores mature, with a small, we did not observe insects near the sporocarps in hyaline stalk, evanescent. Ascospores globose the field. (22 )24 28 ( 30) mm(mean 25.8 mm) includ- ing ornamentation; walls 2 3 mm broad, with light The brown, scruffy peridium and spore size microscopy ornamentation in spore surface view resemble E. austrogranulatus Castellano, Trappe appearing vermiculate, with irregularly shaped & Vernes from Australia. The peridium of ridges and clumps; in cross-sectional view of the E. austrogranulatus is persistently closed and spore the ornamentation appearing as a palisade tougher (more leather-like) and its spores have of fine spines 1.5 3.5 mm tall, not uniformly spaced, outline of spore commonly flattened in an ornamentation of low amorphous warts that places; SEM reveals an ornamentation of spines are easily distinguishable from the spores of fused together in short ridges or small bundles to E. putridus. form a subreticulum or irregular labyrinth of ridges and bundles, spines are often truncate. PARATYPE. BULLER: Murchison, Matakitaki Spore colour grey brown to black in KOH, Bridge Scenic Reserve, M.A. Castellano & brown to dark brown in Melzer’s reagent. R.E. Beever (Trappe 12566), 23 Sept 1992 (PDD 89288, OSC). HOLOTYPE. BULLER, Murchison, Warbeck Sce- nic Reserve, R.E. Beever & M.A. Castellano (Trappe 12572), 24 Sep 1992 (PDD 89289, isotype Acknowledgements OSC). Unfortunately Dr Ross Beever died while still in the process of preparing this manuscript. RE Beever HABIT, HABITAT AND SEASON. Hypogeous becom- acknowledges support from Foundation for Re- ing emergent at maturity, in Nothofagus forests, search, Science and Technology contract CO9309, putatively mycorrhizal with Nothofagus fusca and Lottery Science, and the New Zealand United States N. menziesii; September. Co-operative Science Programme. JM Trappe and MA Castellano received support from National DISTRIBUTION. New Zealand, North Island. Science Foundation Grant BSR 9201421.
ETYMOLOGY. From Latin putrida,rotting,in reference to the carrion-like smell of the mature References ascoma. Blackwell M, Hibbett DS, Taylor JW, Spatafora JW. 2006. Research coordination networks: a phy- logeny for kingdom Fungi (Deep Hypha). REMARKS. Elaphomyces putridus differs from Mycologia 98: 829 837. all other Elaphomyces species in the scanty Castellano MA, Trappe JM 1990. Australasian development of the mycelial crust, the subepi- truffle-like fungi. I. Nomenclatural bibliogra- Sequestrate fungi of New Zealand 433
phy of type descriptions of Basidiomycotina. Lumbsch HT, Huhndorf SM 2007. Outline of Australian Systematic Botany 3: 653 670. Ascomycota 2007. Myconet 13: 1 58. Castellano MA, Trappe JM 1992. Australasian Miller Jr OK, Henkel TW, James TY, Miller SL truffle-like fungi. V. Nomenclatural bibliogra- 2001. Pseudotulostoma, a remarkable new vol- phy of type descriptions of Ascomycotina and vata genus in the Elaphomycetaceae from Zygomycotina. Australian Systematic Botany 5: Guyana. Mycological Research 105: 1268 1272. 631 638. Reynolds HT 2011. Systematics. phylogeography Castellano MA, Trappe JM, Vernes K 2011. Aus- and ecology of Elaphomycetaceae. Ph.D. tralian species of Elaphomyces (Elaphomyceta- Thesis, Duke University, Durham, North ceae, Euriotales, Ascomycota). Australian Carolina. 182 p. Systematic Botany 24: 32 57. Rodway L 1918. Botanical notes. Papers and Pro- Cooke MC 1892. Handbook of Australian fungi. ceedings of the Royal Society of Tasmania 1917: London, Williams and Norgate. 458 p. 105 110. Dodge CW 1929. The higher Plectascales. Annales Trappe JM 1979. Orders, families and genera of hypogeous Ascomycotina (truffles and their Mycologia 27: 145 184. relatives). Mycotaxon 9: 297 340. Index Fungorum 2011. http://www.indexfungorum. Trappe JM, Castellano MA, Malajczuk N 1992. org/Names/Names.asp (accessed 10 June 2011). Australasian truffle-like fungi. II. Index Herbariorum 2011. http://sciweb.nybg.org/ Labyrinthomyces, Dingleya and Reddellomyces science2/IndexHerbariorum.asp (accessed 10 gen. nov. (Ascomycotina). Australian Systema- June 2011). tic Botany 5: 597 611. Kelly KL, Judd DB 1965. The ISCC NBS method Zhang BC, Minter DW. 1989. Elaphomyces of designating colors and a dictionary of color spinoreticulatus sp. nov. with notes on Canadian names. Washington, DC, National Bureau of species of Elaphomyces. Canadian Journal of Standards Circular 553. 158 p. Botany 67: 909 914.