Truffle-like Basidiomycotina of New Zealand: Gallacea, Hysterangium, Phallobata, and Protubera

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1 New Zealand Journal of Botany ISSN: X (Print) (Online) Journal homepage: Truffle-like Basidiomycotina of New Zealand: Gallacea, Hysterangium, Phallobata, and Protubera Michael A. Castellano & Ross E. Beever To cite this article: Michael A. Castellano & Ross E. Beever (1994) Truffle-like Basidiomycotina of New Zealand: Gallacea,Hysterangium,Phallobata, and Protubera, New Zealand Journal of Botany, 32:3, , DOI: / X To link to this article: Published online: 31 Jan Submit your article to this journal Article views: 278 Citing articles: 12 View citing articles Full Terms & Conditions of access and use can be found at

2 New Zealand Journal of Botany, 1994, Vol. 32: X/94/ $2.50/0 9 The Royal Society of New Zealand Truffle-like Basidiomycotina of New Zealand: Gallacea, Hysterangium, Phallobata, and Protubera MICHAEL A. CASTELLANO U.S. Department of Agriculture, Forest Service Pacific Northwest Research Station Forestry Sciences Laboratory 3200 Jefferson Way Corvallis, Oregon 97331, U.S.A. ROSS E. BEEVER Manaaki Whenua - Landcare Research Private Bag Auckland, New Zealand Abstract Three Gallacea species (including G. dingleyae sp. nov., G. eburnea sp. nov.), five Hysterangium species (including H. neotunicatum sp. nov., H. rugisporum sp. nov., H. rupticutis sp. nov., H. youngii sp. nov.), one Phallobata species, and three Protubera species (P. hautuensis sp. nov., P. nothofagi sp. nov., P. parvispora sp. nov.) are recognised in New Zealand. Keys to and descriptions of all taxa are provided. Keywords truffle-like fungi; Basidiomycotina; fungi; biodiversity; systematics; Gallacea; Hysterangiurn; Phallobata; Protubera INTRODUCTION The truffle-like (gastroid) Basidiomycotina comprise those species with basidia that mature in an enclosed basidioma (angiocarpic), with spores that are not forcibly discharged (statismosporic), and in which the basidioma itself is indehiscent. These taxa traditionally have been grouped in the Hymenogastrales, but modern studies have shown that this is a polyphyletic group, and most species B92048 Received 15 December 1992; accepted 10 February 1994 can be placed in well-recognised orders alongside species with dehiscent basidiomata. Basidiomata of many of the truffle-like species are globose to subglobose and superficially resemble each other in the field. Most are hypogeous, producing basidiomata beneath the soil surface, although others, especially the stipitate (secotioid) species, are partially exposed or fully exposed (epigeous). Many form mycorrhizae with shrubs and trees, although definitive evidence is available for relatively few species in New Zealand (Chu-Chou & Grace 1983). The only comprehensive taxonomic treatment of this group in New Zealand is that of Cunningham (1944), but refinement of many generic concepts and availability of additional collections have outdated this account. Although the stipitate species, which Cunningham mostly placed in Secotiurn, have been revised and placed in a variety of genera including Weraroa (Singer & Smith 1958) and Thaxterogaster (Horak 1973), the non-stipitate species still await modern treatment. The present account covers those species included by Cunningham (1944) in Hysterangium and newly recognised congeneric species. We follow recent treatments that place the genera Hysterangium and Protubera in the Phallales, although some authors, while accepting a close evolutionary relationship with the Phallales, segregate them into the Hysterangiales (e.g., Jtilich 1981). Phallobata in our view is clearly allied to Protubera, and thus appropriately placed in the Phallales. We feel less comfortable about placing the less gelatinised, relatively large spored Gallacea in this order, but at this time we suggest no alternative. METHODS Macroscopic features are described from fresh material where collection notes were available. Odour was assessed directly on collection or after enhancement by incubation for a few hours in a sealed vial. Chemical tests involved observing colour reaction of fresh tissues to 95% ethanol (EtOH), 10%

3 306 New Zealand Journal of Botany, 1994, Vol. 32 FeSO4, 5% KOH, and Melzer's reagent. Microscopic features are described from dried material mounted in KOH and in Melzer's reagent and examined by bright field and Nomarski interference microscopy. Material for scanning electron microscopy (SEM) was rehydrated in KOH and processed by ethanol or DMP dehydration and critical point drying essentially as described by Beaton et al. (1985a); photomicrographs were taken on a Philips 505 instrument (mention of a trade name does not constitute endorsement by the USDA). Habitat and fruiting season details refer only to New Zealand. Distributions are based on collections examined by us. Collection details are cited only for New Zealand collections and extralimital types. Associated mycorrhizal host species are indicated where information is available: "Leptospermum spp." is used to refer to Leptospermum scoparium and/or Leptospermum (Kunzea) ericoides, as field notes are often not explicit as to which species was present. New Zealand collection sites are cited according to collecting districts (Crosby et al. 1976). Herbarium names are abbreviated according to Holmgren et al. (1990). Keys are provided to the taxa treated in this paper, but some imperfectly known taxa are not included, and others probably remain to be described. KEY TO GENERA The genera included in this account are characterised by non-stipitate, gelatinised basidiomata lacking a percurrent columella, and by non-amyloid, sometimes slightly dextrinoid, smooth or slightly verrucose, hyaline or pale green to olive brown, oblong to fusiform spores, which may or may not be enclosed by an utricle. 1 Basidiomata convoluted with conspicuous, sterile or partially fertile, gelatinous lobes; spores 2-3.5(-4) lain, utricle absent. Phallobata Basidiomata usually more or less globose, if lobed then with gleba penetrating lobes; spore length 4 [am and width 1.5 pro, utricle present or absent Peridial subcutis thick, gelatinous, transected by sutures connecting epicutis to glebal locules; spore length 7 ~tm and width 3.5 ~tm, utricle absent... Protubera Peridial subcutis not conspicuously gelatinised, lacking sutures; spore length 6 pm and width 3 ~tm, utricle present or absent Gleba brown to olive, often developing large (to 2 cm) schizogenous cavities; spores orthotropic to slightly heterotropic, utricle absent Gallacea Gleba green to olive, not developing schizogenous cavities; spores orthotropic, utricle present... Hysterangium GALLACEA LLOYD, LYCOPERD. AUSTRALIA & NEW ZEAL. P. 37, Basidiomata large (to 10cm), globose to subglobose, broadly obpyriform or irregularly lobed. Peridium thick (c pm), white to pale yellow, pale pink, violet, or purple, bruising pinkish brown to brown, occasionally separable from gleba. Gleba olive to brown, with more or less radially elongate locules partially filled with spores, often developing large (up to 2 cm) schizogenous cavities. Oflour nil to acetylenic or sweetish putrid. Basidia in a euhymenium, hyaline, thin-walled, 4-6(-7)- spored. Subhymenium poorly developed to inconspicuous. Spores statismosporic, orthotropic to slightly heterotropic, smooth, ellipsoid to oblong, lacking an utricle, with a distinct sterigmal attachment, pale green, yellow olive, or pale olive brown, slightly dextrinoid in Melzer's reagent. Development angiocarpic, hypogeous to subepigeous. ETYMOLOGY: From Latin galla, the oak-apple (oak gall), referring to the resemblance of the basidioma to such galls. TYPE SPECIES: Gallacea scleroderma (Cooke) Lloyd. REMARKS: Gallacea was erected by Lloyd to accommodate G. scleroderma, and G. avellana Pat. was subsequently described from New Caledonia (Patouillard 1910). Cunningham (1934) referred G. scleroderma to Hysterangium, but we prefer to retain Gallacea because of the absence of an utricle from the spore and the tendency for large schizogenous cavities to develop in the gleba, and refer two new taxa to the genus. Cunningham (1924) studied the development of G. scleroderma. Key to Gallacea species 1 Basidiomata purple to violet; peridium singlelayered; spores x 4-5 grn; associated with Nothofagus... G. scleroderma Basidiomata white to pale yellow; peridium 2- layered; associated with Nothofagus or Leptospermum Peridium ~tm thick, hyphae of subcutis

4 Castetlano & Beever--Truffie-like fungi 307 A.=._, Fig. 1 Gallacea dingleyae. A, Basidioma exterior and in section (PDD 55927). B, Basidiospores by bright field (PDD 55927). C, Basidiospores by SEM (PDD 55927). Scale: 1 cm (A), 10 lam (B, C). periclinally elongate, interwoven; spores 9-11 x pm; associated with Leptospermum G. dingleyae Peridium pm thick, hyphae of subcutis isodiametric; spores (6-)7-9 x 3-4pm; associated with Nothofagus... G. eburnea Gallacea dingleyae Castellano & Beever sp. nov. (Fig. 1) Basidiomata usque ad 5 cm diam., globosa vel obpyriformia, eburnea, irregulariter avellaneo-columbrina. Gleba fusca vel brunnea, aliquot punctis lacteis. Rhizomorphae evanescentes. Columella dendroidea, crassa, gelatina vel lactitranslucens. Peridium prn crassum; epicute prn crassa, subcute prn crassa. Trama gm crassa. Sporae laeves, 9-11 x 4-5.5/am, oblongae vel ellipsoideae, interdum parum asymmetricae, apice obtuso, base truncata, singulariter pallide olivaceo-bmnneae, aggregate avellaneae. Holotypus: PDD Basidiomata up to 5 cm diam., globose to broadly obpyriform, yellowish white, irregularly mottled pale yellowish brown, surface faintly fibrillose, dull when fresh becoming sheeny on drying, sometimes cracking. Gleba dark greyish brown to moderate brown with occasional milky white flecks, locules more or less radially elongate, partially filled with spores, eventually deliquescing deep brown. Rhizornorphs evanescent. Columella dendroid, stout, with distinct branches to mid-sporocarp, gelatinous to milky translucent, with a nongelatinised, cottony, white region near base. Taste slightly peppery. Odour sweetish putrid when deliquescent. Peridium separable, pm thick, 2- layered; epicutis pm thick, of yellowish

5 308 New Zealand Journal of Botany, 1994, Vol. 32 brown, thin-walled, collapsed, periclinal hyphae 1-3 pm diam., clamp connections absent; subcutis pm thick, of hyaline, thin-walled, interwoven hyphae, (3-)5-10(-25) pm diam., in longitudinal section appearing periclinally elongate with lenses of isodiametric hyphae, clamp connections absent, near gleba a band of collapsed, yellowish brown hyphae overlying sterile locules lined by basidium-like inflated cells to 25 prn diam.; sutures absent. Trama pin thick, of hyaline, collapsed, compactly interwoven hyphae, 1-5(-6) prn diam., in a gelatinised matrix, clamp connections absent. Basidia cylindrical to subclavate, up to 8 pm diam. at apex, c. 5 jam at base, thin-walled, hyaline, (4-)6- spored. Spores smooth, 9-11 x pm, oblong to ellipsoid, sometimes slightly asymmetric, apex obtuse, base truncate, sometimes with a sterigmal attachment pm long by 1.5 ~m wide; wall c. 1 pm thick. Utricle absent. Spore eolour in KOH pale olive brown singly, yellowish brown in mass. Habitat hypogeous or subepigeous in scrubland and modified forest, putatively mycorrhizal with Leptospermum spp. Season April to July. Distribution New Zealand. ETYMOLOGY: Named for New Zealand mycologist Joan M. Dingley, who has collected many trufflelike basidiomycetes. TYPE: NEW ZEALAND, TAUPO, Tongariro National Park, Ketetahi Track, R.E. Beever 872, 19 May 1989 (holotype PDD 55927; isotype OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: AUCKLAND: Waitakere Ranges, Huia, J.M. Dingley, Jul 1948 (PDD 6768, OSC); Waitakere Ranges, Oratia, J.M. Dingley, 2 May 1970 (PDD 28266, OSC)", Waitakere Ranges, Maungaroa Ridge Track, R.E. Beever 963, 21 Apr 1990 (PDD 57240). REMARKS: Gallacea dingleyae is distinguished from G. eburnea by its thinner peridium and larger spores. Gallacea eburnea Castellano & Beever sp. nov. (Fig. 2) Basidiomata usque ad 6 cm diam., globosa, subglobosa vel irregulariter lobata, alba, eburnea vel pallide persicina, contundentibus persicino-brunnea, interdum rimosa. Gleba saturate olivacea vel olivaceo-brunnea, loculis irregularibus vel radiatim elongatis. Rhizomorphae evanescentes, numerosae, parvae, peridio concolorae. Columella dendroidea, gelatina, lactitranslucens. Peridium ~am crassum; epicute wn crassa, subcute 500~600 prn crassa. Trama pm crassa. Sporae laeves (6-)7-9 x 3-4 gm, oblongae, apice obtuso, base pedicellata, singulariter pallide virides, aggregate pallide olivaceo-brunneae. Holotypus PDD Basidiomata up to 6cm diam., globose to subglobose or irregularly lobed, larger sporocarps irregularly grooved and dimpled, white to pale yellow or pale pink when fresh, bruising pinkish brown, drying pale yellow brown to brown, surface glabrous, sometimes cracking to expose gleba; EtOH slowly pale pink, FeSO4 blue green, KOH pale yellow orange, Melzer's reagent negative. Gleba dark olive to olive brown, locules irregular to radially elongate, partially filled, developing schizogenous cavities up to 2cm across. Rhizomorphs evanescent, numerous, branched, attached to indented base, concolorous with peridium. Columella dendroid, with up to 5 primary branches in longitudinal section, gelatinous, milky translucent, base traversed by strands of opaque white tissue arising from peridium. Taste nil to slightly peppery. Odour acetylenic after incubation. Peridinm not separable from gleba, pm thick, 2-layered; epicutis pm thick, of pale brown, thin-walled, somewhat gelatinised, interwoven to subpericlinal hyphae, 1-3 pm diam., clamp connections absent; subcutis pm thick, of hyaline, thin-walled, irregularly shaped isodiametric inflated cells, to ~tm diam., clamp connections absent. Trama pm thick, of hyaline, collapsed, loosely interwoven hyphae, 1-4 lam diam., in a gelatinised matrix, clamp connections absent. Basidia rehydrating poorly, thin-walled, hyaline, 6(- 7)-spored. Spores smooth, (6-)7-9 x 3-4 pro, oblong, sometimes slightly asymmetric; apex obtuse, base with a distinct sterigmal attachment 1 1 pm; wall <0.5 pm thick. Utricle absent. Spore colour in KOH pale green singly, pale olive brown in mass. Habitat hypogeous or subepigeous in Nothofagus forest, putatively mycorrhizal with Nothofagus fusca, N. menziesii, N. solandri, and N. solandri var. cliffortioides. Season February to September. Distribution New Zealand. ETYMOLOGY: From Latin eburneus, in reference to the predominantly white colour of the fresh peridium. TYPE: NEW ZEALAND, TAUPO, Tongariro National Park, Whakapapanui Track, R.E. Beever 711, 8 May 1987 (holotype PDD 48798; isotypes OSC, herb. T. Hongo). OTHER COLLECTIONS EXAMINED: NEW

6 Castellano & Beever--Truffie-like fungi 309 Fig. 2 Gallacea eburnea. A, Basidiomata exterior and in section (PDD 48798). B, Basidiospores by bright field (PDD 48798). C, Basidiospores by SEM (PDD 48798). Scale: 1 cm (A), 10 ~m (B, C). ZEALAND: TAUPO: Tongariro National Park, Whakapapanui Stream, R.F.R. McNabb, 30 Apr 1967 (PDD 45878, OSC); Turangi, M. Chu-Chou, 17 Apr 1980 (NZFR13295); Turangi, M. Chu-Chou, 22 Apr 1981 (NZFRI 3296a); Turangi, M. Chu- Chou, Apr 1982 (NZFRI 3297); Turangi, M. Chu- Chou, 18 May 1983 (NZFRI 3294); Kaimanawa Forest Park, Clements Road, C. Tanaka, 4 Feb 1989 (PDD 56739); Kaimanawa Forest Park, Tree Trunk Gorge, G.M. Taylor, 19 May 1989 (PDD 55924, OSC). NELSON: Abel Tasman National Park, road to Canaan, R.E. Beever 561, 27 Apr 1986 (PDD

7 310 New Zealand Journal of Botany, 1994, Vol. 32 Fig. 3 Gallacea scleroderma. A, Basidioma exterior and in section (PDD 48939). B, Basidiospores by bright field (PDD 48939). C, Basidiospores by SEM (PDD 48936). Scale: I cm (A), 10 jam (B, C) ); Glenhop'e Scenic Reserve, Granity Creek, R.E. Beever 1281, 25 Sep 1992 (PDD 60559). BULLER: Warbeck Scenic Reserve, Maruia Saddle, R.E. Beever211, 16 Apr 1983 (PDD 48792, OSC); same locality, R.E. Beever 545, 24 Apr 1986 (PDD 48800, OSC); same locality, R.E. Beever548, 24 Apr 1986 (PDD 48796, OSC); Nelson Lakes National Park, Lake Rotoroa, R.E. Beever 534, 23 Apr 1986 (PDD 48794, OSC); same locality, R.E. Beever 535, 23 Apr 1986 (PDD 48803, OSC). Victoria Forest Park, Shenandoah Saddle, K. Aoshima & R.E. Beever544, 24 Apr 1986 (PDD 48795, OSC); Buller Gorge Scenic Reserve, R.E. Beever 1275, 23 Sep 1992 (PDD 60561); Murchison, Matakitaki Bridge Scenic Reserve, R.E. Beever 1276 & M.A. Castellano, 24 Sep 1992 (PDD 60560). NORTH CANTERBURY: Arthur's Pass National Park, pass to No. 2 hut, M.J. Noonan, 7 Jun 1971 (PDD 31458, OSC); Craigieburn Forest Park, Cave Stream Shelter, R.E. Beever 521, 19 Apt 1986 (PDD 48799); same locality, R.E. Beever 636, 22 Apr 1987 (PDD 48793, OSC). FIORDLAND: Eglinton Valley, E.E. Chamberlain, 17 May 1963 (PDD 21165, OSC); Te Anau, Kepler Track, R.E. Beever 998, 17 May 1990 (PDD 57833); Lake Gunn, P.R. Johnston, 17 May 1990 (PDD 57832); Lake Hauroko, Lookout Track, R.E. Beever 1030, 21 May 1990 (PDD 57831). REMARKS: Gallacea eburnea differs from G. dingleyae in its smaller spores and thicker peridium. G. avellana Pat. from New Caledonia resembles both these species in its white peridium, but differs in its non-olivaceous reddish gleba and smaller (5-6 x 3 ~tm) spores. Chu-Chou & Grace (1983) illustrate G. eburnea as "Hysterangium species 4".

8 Castellano & Beever Truffle-like fungi 311 Gallacea scleroderma (Cooke) Lloyd, Lycoperd. Australia & New Zeal. p. 38, (Fig. 3) - Mesophellia scleroderma Cooke, Grevillea 14: 11, Hysterangium sclerodermum (Cooke) Cunningham, Proc. Linn. Soc. New S. Wales 59: 165, = Rhizopogon violaceus Cooke & Massee in Cooke, Grevillea 21: 1, = Gallacea violacea (Cooke & Massee) Lloyd, Mycol. Writings 7: 1201, Basidiomata up to 10cm diam., globose to depressed, larger specimens irregularly dimpled and grooved, purple to violet when fresh, paler where exposed, bruising brown, surface tomentose to scaly. Gleba olive brown, deep yellowish brown to dark greyish brown; locules elongate to irregular, partially filled with spores; developing large schizogenous cavities up to 2 cm diam. at maturity. Rhizomorphs attached at base, up to 1 mm diam., concolorous with peridium near base, white distally. Columella often poorly developed, dendroid, translucent when fresh, dark brown when dried. Taste slightly peppery. Odour nil. Peridium not separable from gleba, a single layer, up to 1500/am thick, of hyaline, thin-walled, loosely interwoven, somewhat gelatinised hyphae 7-8/am in diam. near gleba, with inflated, isodiametric cells up to 70/am diam. near surface, clamp connections absent; surface cells yellowish brown with areas of pale violet to violet pigment (when mounted in KOH) and adherent crystalline particles; near gleba sterile locules contiguous with subcutis and lined with basidium-like inflated cells, sometimes with locule wall against gleba fertile. Inflated cells poorly developed in young specimens. Sutures absent. Trama up to 70/am thick, of hyaline, thin-walled, mostly collapsed, subparallel to loosely interwoven hyphae, 2-3/am diam. in a gelatinised matrix, clamp connections absent. Basidia elongate to subclavate, 6-8/am diam. at apex c. 3/am at base x 24-35/am, hyaline, thin-walled, 4-6-spored. Spores smooth, /am, ellipsoid; apex obtuse, base obtuse, with or without a distinct asymmetric sterigmal attachment up to 2/am long 1 pm wide; wall c. 1 prn thick. Utriele absent. Spore eolour in KOH yellow olive singly, brown in mass. Habitat hypogeous or subepigeous in Nothofagus forests, putatively mycorrhizal with Nothofagus fusca, N. menziesii, N. solandri, and N. solandri var. cliffortioides. Season January through October. Distribution New Zealand. ETYMOLOGY: Literally from Greek sclero, hard, and derma, skin, but in this context in reference to the macroscopic resemblance of the basidioma to those of Scleroderma spp. TYPE: NEW ZEALAND, locality unknown, J. Reader 50, Apr 1885 (holotype K). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: WELLINGTON: York Bay, E.H. Atkinson, 5 Oct 1919 (PDD 839, OSC); same locality, E.H. Atkinson, 4 Feb 1922 (PDD 823); same locality, E.H. Atkinson, 26 Jun 1922 (PDD 703, OSC, BPI); same locality, E.H. Atkinson, 10 Oct 1922 (PDD 926, OSC); same locality, E.J. Butler & G.H. Cunningham, 27 Jul 1923 (PDD 1220, OSC). NELSON: Nelson City, track to Fringed Hill, R.E. Beever 666, 30 Apr 1987 (PDD 48939, OSC). BULLER: Murchison, Matakitaki Bridge Scenic Reserve, P.K. Buchanan & R.E. Beever647, 27 Apr 1987 (PDD 48938, OSC); same locality, R.E. Beever 648, 27 Apr 1987 (PDD 48937); same locality, R.E. Beever 1272, 23 Sep 1992 (PDD 60558); same locality, R.E. Beever 1277, 24 Sep 1992 (PDD 60557); Nelson Lakes National Park, Lake Rotoiti, Loop Track, R.E. Beever 659, 28 Apr 1987 (PDD 48940, OSC). NORTH CANTERBURY: Arthur's Pass, R.G. MacDougall, 27 May 1960 (PDD 19314, OSC); Hawdon Valley, Woolshed Hill, S.J. Hughes 720, 16 May 1963 (DAOM 93583, OSC); Cass, M.J. Noonan, 30 Mar 1968 (PDD 31459, OSC); Lewis Pass, B.S. Parris, 26 Jan 1970 (PDD 28165, OSC); Arthur's Pass National Park, Casey Saddle, M.J. Noonan, 4 Jun 1972 (PDD 31390, OSC). MID CANTERBURY: Upper Waimakariri, T. Kirk 382,?date, (type of Rhizopogon violaceus, holotype K); Upper Rangitata River, V. Begg, 27 May 1961 (PDD 19830, OSC); Craigieburn Forest Park, Cave Stream Shelter, R.E. Beever515, 19 Apr 1986 (PDD 48941); same locality, R.E. Beever 629, 22 Apr 1987 (PDD 48942). OTAGO LAKES: Queenstown, One Mile Creek, R.E. Beever276, 15 May 1984 (PDD 48936); Lake Hawea, Hunter Valley Road, Sawyer Burn, A. Newton & M.K. Thayer, 17 Jan 1985 (PDD 48943). FIORDLAND: Fiordland National Park, Waiau River, Kepler Track, R.E. Beever 1012, 18 May 1990 (PDD 57834). REMARKS: Gallacea scleroderma, a common species in Nothofagus forests, is readily recognised in the field by the purple pigmentation of its peridium. One collection [WELLINGTON, Tararua Range, Mt Reeves, T.C. Birch, Mar 1931 (PDD 8316, DAOM , OSC)] determined as G. scleroderma on the basis of peridial characters,

9 312 New Zealand Journal of Botany, 1994, Vol. 32 including traces of purple pigmentation, approaches G. eburnea in spore size (6-7 x pm). HYSTERANGIUM VITTADINI, MONOGR. TUBERAC. P. 13, Basidiomata globose to subglobose or irregular. Peridium to 1 mm thick, white to salmon pink, pale yellow, or pale brown, often staining pink to brown where bruised, sometimes readily separable from gleba, bearing hyphae and rhizomorphs at base and sides or overall. Gleba grey, greyish green or olive, with small labyrinthine locules and a poorly to strongly developed, dendroid, gelatinous to cartilaginous, white or grey columella. Basidia in a euhymenium, hyaline to occasionally pale brown, thin-walled, 2-6-spored. Subhymenial layer poorly developed. Spores statismosporic, orthotropic, smooth to moderately verrucose, with a wrinkled to loose utricle, narrowly ellipsoid to fusoid, sterigmal attachment straight, inconspicuous to truncatecupped, hyaline, pale green or pale olive singly, slightly dextrinoid in Melzer's reagent. Development angiocarpic, hypogeous to subepigeous. ETYMOLOGY: From Greek hystera, womb, and angeion, vessel (a term used by 19th century mycologists for a sporocarp), referring to the angiocarpic basidioma. TYPE SPECIES: Hysterangium clathroides Vittadini, designated by Zeller & Dodge (1929). REMARKS: Hysterangium is a large genus, widely distributed in both Northern and Southern Hemispheres, forming ectomycorrhizae particularly with Pinaceae, Fagaceae, and Myrtaceae. In the following descriptions, the ornamentation of the spore wall is described as it appears by light microscopy, although in some species the utricle is appressed regularly to the spore wall, preventing precise elucidation of the wall details. Key to Hysterangium species 1 Spores 18 ~tm long, moderately verrucose, utricle adhering, seldom inflated... H. youngii Spores <18 ~tm long, minutely verrucose or smooth, utricle various Peridium of interwoven, sometimes inflated hyphae; spores smooth, 9-12 x 4-5 lain, utricle distinctly inflated... H. inflatum Peridium with isodiametric cells; spores smooth or verrucose, >12 ~tm long, utricle various Peridium 3-layered; epicutis of interwoven beadlike hyphae; spores verrucose, gm long, utricle adhering to spore wall... H. rugisporum Peridium 2-layered; epicutis of isodiametric cells, subcutis of interwoven hyphae; utricle adhering to spore wall or inflated Spores smooth, ~tm long, wall 0.5 gm thick, utricle irregularly inflated; associated with Leptospermum... H. neotunicatum Spores verrucose, ~tm long, wall c. 1 ~tm thick, utricle usually adhering to spore wall; associated with Eucalyptus... H. ruptieutis Hysterangium inflatum Rodway, Paps. Proc. Roy. Soc. Tasmania 1917: 109, (Fig. 4) = Hysterangium eucalyptorum Lloyd, Mycol. Writ. 7:1119, = I4ysterangiumpterosporum Donadini & Riousset, Trav. Sci. Parc Nation. PortCros 5:11, Basidiomata up to 2.5 cm diam., globose to subglobose, ellipsoid or irregularly lobed, dull white with pale brown to pale reddish brown patches when fresh, drying dark brown except in recessed areas which are pale brown to almost white, surface smooth to mostly tomentose when fresh, wrinkled when dry, surface with adhering soil particles. Gleba olive grey to bright green when young, dark olive to almost black when mature and deliquescent; locules more or less radially elongate, partially to completely filled with spores. Rhizomorphs numerous, highly branched, attached at base, concolorous with peridium, associated with conspicuous white mycelium in soil. Columella dendroid, narrow, translucent. Taste nil to faintly peppery. Odour slightly perfumed to sharp after incubation. Peridium separable from gleba in large flakes, a single layer (-250) ~rn thick, of brown (near surface) or hyaline (near gleba), thin-walled, irregularly shaped, sometimes inflated hyphae, cells elongate to subglobose, up to 5 ~tm diam. near gleba, up to 12 x 25 tam diam. in mediostratum, with numerous minute crystals adhering to outer hyphae, clamp connections common. Trama lam thick, of hyaline, loosely to compactly interwoven hyphae, ~tm diam., in a gelatinised matrix, hyphae mostly elongate, sometimes with nodose ends, clamp connections absent; with suture-like structures traversing pefidial trama in places. Basidia rehydrating poorly, hyaline, (4-)6-spored. Spores smooth, 9-12 x (3-)4-5 ~tm without utricle, 9-12 x 8-9 ~tm with utricle, ellipsoid; apex

10 Castellano & Beever--Truffie-like fungi 313 Fig. 4 Hysterangium inflatum. A, Basidioma in section and exterior (PDD 48309). B, Basidiospores by bright field (Trappe 9905, OSC). C, Basidiospores by interference contrast (Trappe 9905, OSC). D, Basidiospores by SEM (PDD 48309). Scale: 1 cm (A), 10 [am (B, C), 5 [am(d). obtuse, base sessile, sometimes truncate; wall <0.5 lam thick. Utricle distinct, laterally inflated up to 2.5 ~tm, attached near base and below apex, commonly forming a cylinder around spore. Spore eolour in KOH hyaline singly, pale green in mass. Habitat hypogeous or subepigeous in plantations and modified communities, putatively mycorrhizal with Eucalyptus spp. including E. delegatensis, E. fastigata, E. nitens, E. regnans (Chu-Chou & Grace 1983). Season April to September. Distribution Algeria, Australia, Ecuador, France, Italy, New Zealand, U.S.A. (California). ETYMOLOGY: From Latin inflatus, swollen, referring to the greatly inflated spore utricle. TYPE: AUSTRALIA, Tasmania, Mt Wellington, Feb 1904, L. Rodway 1267 (holotype HO 89531; isotypes OSC, NY, FH). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: COROMANDEL: Thames, Waiomu Valley, R.E. Beever 748, 29 Apr 1988 (PDD 48309); same locality and date, M.P. Amaranthus (Trappe 9904, 9905, 9906), (OSC), M.A. Castellano (Trappe 9895, 9897, 9898, 9903), (OSC), R. Young (Trappe 9887, 9899, 9900, 9901, 9902), (OSC). TAUPO: Tokoroa, M. Chu-Chou, 5 Apr 1979 (NZFRI 3287); Kaingaroa Forest, M. Chu-Chou, May 1982 (NZFRI 3289). TARANAKI: Egmont National Park, Kaitake Range, Lucys Gully, R.E. Beever687, 16 May 1987

11 314 New Zealand Journal of Botany, 1994, Vol. 32 (PDD 48310, OSC). MID CANTERBURY: Christchurch, Victoria Park, R.E. Beever 1265, 20 Sep 1992 (PDD 60576). ECUADOR: Quito, L. Mille, Jan 1921 (type ofh. eucal~ptorum, NY, PDD 8309). FRANCE: Porquerolles, L. Riousset, 12 Nov 1980 (OSC, det. by Riousset as H. pterosporum). REMARKS: The description includes data from the many Australian collections of the first author. Hysterangium inflatum is easily distinguished from all other species by its regularly inflated, barrel-like utricle, which gives the spore a winged appearance in optical section. Hysterangium inflatum appears to be a common ectomycorrhizal associate of Eucalyptus spp. and has evidently been introduced with Eucalyptus seedlings to many parts of the world. In contrast to Beaton et al. (1985b), we regard Hysterangium tunicatum Cunningham as a distinct, albeit invalidly published, species. H. pterospermum is placed in synonymy after examining an authenticated collection. The species was first recorded from New Zealand by Chu-Chou & Grace (1983). Their "Hysterangium sp. 3" [?locality, M. Chu-Chou, 6 Jul 1979 (NZFRI 3293)] appears also to be this species. The Zeller & Dodge (1929) record of H. inflatum from New Zealand was based on Cunningham collection "1189" (PDD 8318) now recognised as H. neotunicatum. Hysterangium neotunicatum Castellano & Beever sp. nov. =Hysterangium tunicatum Cunningham, Trans. Roy. Soc. New Zeal. 67: 409, nora. nudum (Fig. 5) Basidiomata usque ad 2.5 cm diam., globosa, subglobosa, irregularia vel reniformia, albida, avellanea vel viridi-grisea, contundentibus pallide rufa, rufa vel cinereae. Gleba cinereoolivacea, olivacea vel obscure cinereo-olivacea; loculis elongatis. Rhizomorphae singulae, crassae, usque ad 1 mm diam., peridio concolorae. Columella dendroidea, gelatina, opaca vel translucens. Sapor parum piperatus. Odor mucido-suavis vel acetylenicus. Peridium vivum facile separabile, Jam crassum; epicute pan crassa, cellulis inflatis lam diam.; subcute ~tm crassa, hyphis intertextis 3-8 gm diam. Trama 50-70(-200) lam crassa. Sporae laeves, x (4-)5-6.5 ~tm, fusiformes vel late fusiformes, apice acuminato; utriculus distinctus, lateraliter inflatus usque ad 2 grn, irregulariter laxus; singulariter pallide virides, aggregate virides. Holotypus PDD Basidiomata up to 2.5 cm diam., globose to subglobose, irregular or reniform, when fresh dull white to yellowish brown or greenish grey and sometimes cracked where exposed, bruising pale reddish brown to reddish brown or grey, surface somewhat glabrous; EtOH pale pink to red. Gleba olive to dark greyish olive; locules elongate, empty or partially filled, deliquescing. Rhizomorph single, stout, up to 1 mm diam., soon branched, attached to slightly indented base, concolorous with peridium. ColumeUa dendroid, gelatinous, opaque to translucent, arising from a sterile base. Taste slightly peppery. Odour musty sweet to slightly acetylenic, becoming sweet-smelling on deliquescing. Peridium easily separable from gleba, ~tm thick, 2-layered; epicutis grn thick, of hyaline, thin-walled, irregularly inflated cells (parenchyma-like), lam diam., smaller near outer surface, clamp connections absent; subcutis am thick, of hyaline, thin-walled, interwoven hyphae, 3-8 ~tm diam., clamp connections absent. Trama 50-70(-200) lam thick, of hyaline, compactly interwoven hyphae, 2-3/am diam., in a gelatinised matrix, clamp connections absent. Basidia rehydrating poorly, hyaline, 6-spored. Spores smooth, x (4-)5-6.5 lam, fusiform to broadly fusiform, apex acuminate, base with a short sterigmal attachment; wall <0.5 ~tm thick. Utriele distinct, laterally inflated up to 2 lam, irregularly loose or baggy. Spore colour in KOH pale green singly, green in mass. Habitat hypogeous to subepigeous in scrubland and modified forest, putatively mycorrhizal with Leptospermum ericoides and L. scoparium. Season throughout the year. Distribution Argentina, Australia, New Zealand. ETYMOLOGY: From Greek neos, new, and tunicatum, in reference to the invalidly published earlier name, which presumably referred to the tuniclike inflated utricle. TYPE: NEW ZEALAND, TAUPO, Tongariro National Park, Ketetahi Track, R.E. Beever 873, 19 May 1989 (holotype PDD 55923; isotype OSC); same locality and date, R.E. Beever 875, 876 (paratypes PDD 55925, 55926, OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: NORTHLAND: Russell State Forest, Punaruku, V.C. Haydon, 12 Apr 1974 (PDD 32186); Whangaruru Harbour, Oakura Bay, V.C. Haydon, 9 Jul 1974 (PDD 32293); Bay of Islands, Moturoa Island, R.E. Beever 1056, 25 Aug 1990 (PDD 57836); same locality, R.E. Beever 1058, 26 Aug 1990 (PDD 57843); same locality, R.E. Beever 1063, 27 Aug 1990 (PDD 57841); same locality, R.E. Beever 1067, 29 Aug 1990 (PDD 57839). AUCKLAND: Kawau Island, J.C. NeiU, Aug 1943

12 Castellano & Beever--Tmffle-like fungi 315 Fig. 5 Hysterangium neotunicatum. A, Basidioma exterior and in section (PDD 51927). B, Detail of basidioma attachment region in section (PDD 51927). C, Basidiospores by bright field (PDD 55923). D, Basidiospores by SEM (PDD 8321, type ofh. tunicatum). E, Basidiospores by SEM (PDD 55923). Scale: 1 cm (A), 1 mm (B), 10 ~m (C, D, E).

13 316 New Zealand Journal of Botany, 1994, Vol. 32 (PDD 8320); Woodhill, J.M. Dingley, 19 Jul 1953 (PDD 12226); Waitakere Ranges, Karekare, R.F.R. McNabb, 9 Jun 1965 (PDD 24551, OSC); Waitakere Ranges, Piha Valley, J.M. Dingley, 20 Jun 1970 (PDD 28379); same locality and collector, 18 Sep 1970 (PDD 28579, OSC); same locality and collector, 26 May 1971 (PDD 29232); Rangitoto Island, J.M. Dingley, 2 Jun 1972 (PDD 30171); Kaipara Harbour, South Head, Coal Seam Ecological Area, R.E. Beever 729, 21 Nov 1987 (PDD 48317, OSC); Waitakere Ranges, Piha Valley Track, R.E. Beever 964, 21 Apr 1990 (PDD 57238). COROMANDEL: Little Barrier Island, J.M. Dingley, 29 Aug 1958 (PDD 20042, OSC); same details (PDD 20041); same details (PDD 20044); same locality, R.E. Beever 344 & E.P. Laracy, 16 Jun 1984 (PDD 48315); same locality, P.K. Buchanan & R.E. Beever 318, 14 Jun 1984 (PDD 48318); Thames, Waiomu Stream, R.E. Beever 599, 7 Aug 1986 (PDD 51927, OSC); same locality, J.R. May, 31 Aug 1986 (PDD 51796); same locality, R.E. Beever 746, 29 Apr 1988 (PDD 48314, OSC); same locality and date, R.E. Beever 747 (PDD 48312); same locality and date, M.P. Amaranthus (Trappe 9883) (OSC); same locality and date, J. Trappe 9885 (OSC); same locality and date, M.A. Castellano (Trappe 9883) (OSC). WAIKATO: Te Aroha, G.H. Cunningham "1189"', 14 Jun 1923 (PDD 8318); Pukemokemoke Hill, R.E. Beever439, 19 Nov 1984 (PDD 48320, OSC). BAY OF PLENTY: Whale Island, R.E. Beever 492, 3 Jan 1986 (PDD 48319, OSC); same locality and date, R.E. Beever493 (PDD 48316, OSC). TAUPO: Turangi, M. Chu-Chou, 17 Apt 1980 (NZFRI 3301). WELLINGTON: Hokio, G. Cone, 27 Jul 1951 (PDD 10842). NELSON: Fringe Hill, G.H. Cunningham, Nov 1923 (type of H. tunicatum G.H. Cunningham, holotype PDD 8321; isotypes DAOM , OSC). MID CANTERBURY: Banks Peninsula, Okuti Valley Scenic Reserve, R.E. Beever 433 & J.D. Lovis, 14 Oct 1984 (PDD 48313). OTAGO: Dunedin, Whare Flat, J.R. Moore, Feb 1937 (PDD 8322). REMARKS: Hysterangium tunicatum Cunningham was published after 1935 without a Latin diagnosis, so is a nomen nudum in accordance with the Botanical Code, Article The epithet neotunicatum is proposed to accommodate this species, which has spores similar to those of H. inflatum but with an irregularly inflated utricle; the two species differ vastly in peridial characters. Chu- Chou & Grace (1983) illustrate this species as "Hysterangium sp 5", although as their fig. 15 rather than fig. 14. Hysterangium neotunicatum differs from H. rugisporum, which also occurs under Leptospermum, by its smooth spores, 2-layered peridium, and pink to red reaction with EtOH and bruising. The spores of H. neomnicatum are smaller than those of H. youngii. Two collections [AUCKLAND, Waitakere Ranges, Huia Dam Track, R.E. Beever 761, 18 Jun 1988 (PDD 48321, OSC);?locality, M. Chu-Chou, 25 May 1982 (NZFRI 3299)] placed in H. neotunicatum on the basis of peridial characters approach H. rugisporum in spore ornamentation. Hysterangium rugisporum Castellano & Beever sp. nov. (Fig. 6) Basidiomata usque ad 3 cm diam., subglobosa vel irregulariter lobata, basi indentata, alba, avellanea-columbrina, contundentibus brunneolis, minute pubescenti. Gleba bruneolo-grisea vel griseoolivacea vel olivacea; loculis irregularibus vel elongatis, plus minusve radiantibus. Rhizomorpha singularis, usque ad 0.5 mm diam., peridio concolor. Columella dendroidea, crassa, in sicco nigrescens. Peridium in squamis grandibus, paulatim deponens, usque ad 800/am crassum; epicute pm crassa, hyphis hyalinis, compactis, moniliformibus, 2-5(-8) ram diam., crystallis minutis incrustata, fibulae destitutae; mesocute fan1 crassa, hyphis subglobosis vel polybedricis, inflatis, usque ad 20 pm diam.; subcute 60-80(-100) lain crassa, cellulis collapsis, brunneolis. Trama ~m crassa. Sporae minute verrucosae, om, late ellipsoideae, apice acuminato; paries plus minusve 0.5 ~m crassus; utrieulus distinctus, 1 jam crassus, rugosus, plerumque ad parietem sporae adhaerens; sporae singulariter pallide virides, aggregate olivaeae. Holotypus PDD Basidiomata up to 3 cm diam., subglobose to irregularly lobed with an indented base, white when fresh, bruising brown, mottled yellowish brown, white to brown when dry, surface with a fine pubescence which is easily rubbed off on handling, sometimes with adhering soil particles; EtOH mid brown. Gleba brownish grey when fresh drying to greyish olive to olive; locules irregular to elongate, more or less radially arranged, empty. Rhizomorph single, becoming branched several millimetres from point of attachment, up to 0.5 mm diam., attached at base, concolorous with peridium. Columella dendroid, with up to 6 primary branches in longitudinal section, gelatinous, translucent to opaque near base, stout, arising from a sterile base, nearly black when dry. Taste slightly peppery. Odour musty to faintly acidic. Peridium peeling readily in large flakes, up to 800 lam thick, 3-layered; epicutis ~m thick, of hyaline, thin-walled, compactly interwoven, beadlike hyphae, which are elongate to subglobose, 2-5(-8) lam diam., associated with droplets ofoil which become minute crystalline incrustations on storage, clamp connections absent; mesocutis lam

14 Castellano & Beever--Truffle-like fungi 317 Fig. 6 14ysterangium rugisporum. A, Basidiomata exterior and in section (PDD 62455). B, Basidioma in section (PDD 62455). C, Basidiospores by bright field (PDD 4605). D, E, Basidiospores by SEM (PDD 4605). Scale: 1 cm (A, B), 10 vm (C, D, E). thick, of hyaline, thin-walled, compact, irregularly inflated, subglobose to polyhedral hyphae up to 20 jam diam., clamp connections absent; subcutis 60-80(- 100) ~rrt thick, of golden brown isodiametric cells which collapse and then appear as more or less periclinal hyphae, clamp connections absent. Trama ~rn thick, of hyaline, compactly interwoven hyphae, 1-2 ~m diam., in a gelatinised matrix, clamp connections absent. Basidia hyaline, cylindric with narrowed base, ~am long, 4-6-spored. Spores minutely verrucose (by light microscopy), lam, broadly ellipsoid, apex obtuse, base with a short sterigmal attachment; wall c. 0.5 ~m thick. Utriele distinct, 1 ~m thick, wrinkled, usually adhering to spore wall. Spore eolour in KOH pale green singly, greenish brown in mass. Habitat hypogeous to partially epigeous in modified forest, putatively mycorrhizal with Leptospermum scoparium and L. ericoides. Season April through October. Distribution New Zealand.

15 318 New Zealand Journal of Botany, 1994, Vol. 32 ETYMOLOGY: From Latin ruga, wrinkle, and spora, spore, referring to the wrinkled utricle surrounding the spores. TYPE: NEW ZEALAND, COROMANDEL, Little Barrier Island, J.M. Dingley, 9 Oct 1945 (holotype PDD 4605; isotypes DAOM , OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: AUCKLAND, Titirangi, Atkinson Park, R.F.R. McNabb, 11 Jun 1965 (PDD 24592, OSC); same locality, R.F.R. McNabb, 10 Sep 1965 (PDD 24704, OSC); Waitakere Ranges, Piha Valley, J.M. Dingley, 11 Jun 1971 (PDD 29233, OSC); Waitakere Ranges, Goldies Bush, R.E. Beever 1346, 19 June 1993 (PDD 62455, OSC); same locality, R.E. Beever 1371, 10 Jul 1993 (PDD 62454, OSC). COROMANDEL, Moehau Range, Stony Bay, R.E. Beever 1373, 8 Sep 1993 (PDD 62456). REMARKS: Hysterangium rugisporum differs from H. youngii in its smaller spores, and from H. neotunicatum in its regularly verrucose spores, and brown reaction with EtOH and to bruising. Hysterangium rupticutis Castellano & Beever sp. nov. (Fig. 7) Basidiomata usque ad 2.5 cm diam., depresse globosa vel irregularia, sordide alba, contundentibus rosea, glabra, in patellis irregularibus rupta. Gleba obscure glauca, loculis plerumque elongatis, plus minusve radiantibus, partialiter repletis. Rhizomorpha singularis, crassa, plus minusve 1 mm diam., peridio concolor. Columella dendroidea, gelatina, alba, interdum inconspicua. Odor fragrans vel mucidus. Peridium non separabile, /am crassum; epicute 400~J~50 lain crassa, cellulis irregularibus, inflatis usque ad 50 ~am diam., hyphis laxe intertextis tecta; subcute lam crassa, hyphis hyalinis, aliquantum agglutinatis, compacte intertextis, irregularibus, 2-3(- 7.5) Jam diam. Trama Jam crassa. Basidia hyalina, elongata, aliquantum ventricosa, 4-spora, x 6-8/am. Sporae minute verrucosae, x 5-6/am, ellipsoideae vel late ellipsoideae, apice acuto vel aliquando subpapillato; paries 1 ~tm crassus; utriculus distinctus, rugosus, plus minusve ad parietem sporae adhaerens, aliquando usque ad 1 mm inflatus; singulariter hyalinae, aggregatae pallide olivaceae. Holotypus PDD Basidiomata up to 2.5 cm diam., depressed globose to irregularly shaped, dingy white when fresh, bruising pink, grey where exposed, glabrous, exposed portion cracked into irregular plates. Gleba dark greenish grey when fresh, dark olive when dried; locules more or less radially arranged, sometimes lamellate, usually elongate, partially filled. Rhizomorph single, stout, c. 1 mm diam., soon branched, concolorous with peridium. Columella dendroid, gelatinous, white, not well developed, sometimes inconspicuous. Taste not noted. Odour perfumed to musty. Peridium not separable from gleba, 2-layered, lam thick; epicutis 400~450 lam thick, of hyaline, thin-walled, irregularly shaped, inflated cells up to 50 lam diam., cells becoming gradually smaller and pale brown near subcutis, clamp connections absent, epicutis overlain with a thin (<10 lam), loosely interwoven layer of crystalencrusted hyphae 2-3 jam diam.; subcutis }am thick, of hyaline, thin-walled, somewhat agglutinated, compactly interwoven, irregularly shaped hyphae 2-3(-7.5) jam diam., clamp connections absent. Trama ~am thick, of hyaline, thin-walled, compactly interwoven hyphae, 2-3 lain diam., in a gelatinised matrix, clamp connections absent. Basidia hyaline, elongate with a constricted base, somewhat swollen centrally, x 6-8 lam, 4- spored, clamp connections common. Spores minutely verrucose, x 5-6 jam, ellipsoid to broadly ellipsoid, apex acute to occasionally subpapillate, base with a slight sterigmal attachment; wall c. 1 ~am thick. Utriele distinct, wrinkled, more or less adhering to spore wall, occasionally inflated to 1/am. Spore eolour in KOH hyaline singly, pale olive in mass. Habitat hypogeous to subepigeous in modified communities, putatively mycorrhizal with Eucalyptus spp. including E. fastigiata. Season July. Distribution New Zealand. ETYMOLOGY: From Latin ruptus, ruptured, and curls, skin, referring to the cracked peridium of exposed specimens. TYPE: NEW ZEALAND, AUCKLAND, Hunua Ranges, Orere Point, R.E. Beever 228 & G.R. Beever, 2 Jul 1983 (holotype PDD 48322, isotype OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: BAY OF PLENTY: Rotoehu Forest, M. Chu-Chou, 4 Jul 1979 (NZFRI 3292). REMARKS: Known only from New Zealand, but the host association suggests that the species has been introduced from Australia. Chu-Chou & Grace (1983) illustrate this species as "Hysterangium sp 2". Hysterangium youngii Castellano & Beever sp. nov. (Fig. 8) Basidiomata usque ad 1 cm diam., subglobosa vel irregulariter lobata, maxime firm& alba, contundentibus brunnea, minime rugosa. Gleba firma, cartilaginea, obscure viridis, loculis irregularibus, partialiter repletis. Rhizomorphae paucae, basi affixae, peridio concolores. Columella multo deminuta, dendroidea, translucens. Peridium non separabile, 400M50 lam crassum, hyphis irregulariter inflatis, 20-35(-40) ~tm diam., prope

16 Castellano & Beever--Truffle-like fungi 319 Fig. 7 Hysterangium rupticutis A, Basidiomata exterior and in section (PDD 48322). B, Basidium and spores by bright field (PDD 48322). C, Basidiospores by bright field (PDD 48322). D, Basidiospores by SEM (PDD 48322). Scale: 1 cm (A), 10 lain (B, C, D). paginam aureo-brunneis, prope glebam hyalinis. Trama usque ad 80 I.tm crassa. Sporae minute vel moderate verrucosae, 18-20(- 22) x 6-7 ~m, ellipsoideae vel fusiformes, aliquot asymmetricae, apice acuminato vel pro parte maxima papillato; paries lain crassus; utriculus rugosus, usque ad 1 tam crassus; singulariter pallide virides, aggregate olivaceo-brunneae. Holotypus PDD Basidiomata up to 1 cm diam., subglobose to irregularly lobed, white, bruising brown when fresh, dark brown with pale brownish white areas when dried, surface wrinkled when fresh. Gleba firm, cartilaginous, dark green; locules irregular, partially filled, some contiguous with peridium. Rhizomorphs few, small (<1 mm diam.), attached to base, concolorous with peridium. Columella much reduced, dendroid, translucent. Taste not noted. Odour nil. Peridium not separable from gleba, a single layer ram thick, of thin-walled, irregularly

17 320 New Zealand Journal of Botany, 1994, Vol. 32 Fig. 8 Hysterangium youngii. A, Basidiomata exterior and in section (PDD 60093, Trappe 9878 (OSC)). B, Basidiospores in optical section by bright field (PDD 60093). C, Basidiospores in surface view by bright field (PDD 60093). D, Basidiospore by SEM (PDD 60093). Scale: 1 cm (A), 10 pm (B, C, D). inflated hyphae, 20-35(-40) pm diam., smaller and more or less periclinal near gleba and peridial surface, outer portion ( pm) golden brown and encrusted with soil particles, inner portion ( pm) hyaline without incrustations, clamp connections absent. Trama up to 80 lam thick, of hyaline, thin-walled, interwoven hyphae, 3-4 gm diam., in a gelatinised matrix, clamp connections absent. Basidia not observed. Spores minutely to moderately verrucose, 18-20(-22) x 6-7 lam, ellipsoid to fusoid, some asymmetric; apex acuminate to mostly papillate, base with a sterigmal attachment 1 pm long x 2 pm wide; wall lam thick, thicker at spore apex. Utricle wrinkled, adherent, up to 1 jam thick. Spore colonr in KOH pale green singly, olive brown in mass. Habitat hypogeous in podocarp-broadleaf forest, probable mycorrhizal associates uncertain. Season April. Distribution New Zealand. ETYMOLOGY: Named for the collector of the type, Roy Young of Grants Pass, Oregon. TYPE: NEW ZEALAND, COROMANDEL, Thames, Kaitarakihi Track, R. Young (Trappe 9878), 28 Apr 1988 (holotype PDD 60093; isotype OSC). REMARKS: Known only from the type collection. Hysterangium youngii has the largest spores of any Hysterangium species in New Zealand. It resembles H. salmonaceum Beaton, Pegler & Young from Australia in spore size and shape, but can be distinguished by its ornamented spores. PHALLOBATA CUNNINGHAM, TRANS. NEW ZEAL. INST. 56: 73, Basidiomata variously shaped, convoluted and lobed. Peridium white to pale grey, glabrous, thick and gelatinised, not separable from gleba. Gleba green to olive, with elongate to irregular locules filled with spores, not extending into all lobes. Basidia hyaline, 8-spored. Spores statismosporic, orthotropic, smooth, lacking an utricle, ellipsoid, sterigmal attachment inconspicuous, hyaline in KOH singly. No distinctive reaction to Melzer's reagent. Development angiocarpic, epigeous. ETYMOLOGY: From Phallus, the stinkhorn genus, in reference to the phalloid-like spores and basidia, and from Latin lobatus, lobed, referring to the lobed structure of the basidioma. TYPE SPECIES: Phallobata alba Cunningham. REMARKS: Monotypic, known only from New Zealand. The absence of sutures and the small spores

18 Castellano & Beever--Truffle-like fungi 321 Fig. 9 Phallobata alba. A, Basidioma exterior with portion of lobe sliced to expose gleba (PDD 47823). B, Basidioma sliced through glebal region and sterile lobes (PDD 56745). C, Basidiospores by bright field (PDD 1187 ). D, Basidiospores by SEM (PDD 1187). Scale: 1 cm (A, B), 10 ~tm (C); 1 ~tm (D). place this genus in the Phallales near Gelopellis rather than Protubera. The very gelatinous nature of the species makes peridial details difficult to discern in KOH mounts of heat-dried specimens. PhalIobata alba Cunningham, Trans. New Zeal. Inst. 56: 73, Hysterangium lobatum Cunningham, Trans. Roy. Soc. New Zeal. 67: 408, (Fig. 9) Basidiomata to 4.5 cm diam., variously shaped, convoluted and lobed, white to pale grey when fresh, yellow brown when dried, glabrous. Gleba green to olive, gelatinous, some lobes completely sterile; locules elongate to irregular, filled with spores. Rhizomorphs attached at base, up to 1 mm diam., concolorous with peridium. Columella prominent, dendroid, opaque when fresh. Taste not noted. Odour nil. Peridium not separable from gleba, mm thick, 2-layered; epicutis ~tm thick, of hyaline, thin-walled, loosely interwoven hyphae, 4-5 }am diam., with scattered (much more numerous near peridial surface), isodiametric, inflated cells, ~m diam., clamp connections common; subcutis (-5000) ~tm thick, of hyaline,

19 322 New Zealand Journal of Botany, 1994, Vol. 32 thin-walled, loosely interwoven hyphae, 1-2 lam diam., in a gelatinised matrix, clamp connections absent. Sutures absent. Trama up to 50 tarn thick, of hyaline, thin-walled, subparallel to loosely interwoven hyphae, 1-2 lam diam., in a gelatinised matrix, clamp connections absent. Basidia "8-spored" (Cunningham). Spores smooth, 2-3.5(-4) x lam, ellipsoid, apex and base obtuse; wall <0.5 lam thick. Utriele absent. Spore colonr in KOH hyaline singly, pale green in mass. Habitat epigeous on ground and rotting wood in forest. Season December, January, and June. Distribution New Zealand. ETYMOLOGY: From Latin albus, white, referring to the colour of the fresh peridium. TYPE: NEW ZEALAND, WELLINGTON, Paekakariki, Whakatikei Forest Reserve, J.C. Neill & J.G. Myers, 4 Jun 1923 (holotype PDD 1187; isotypes DAOM , FH, K, NY, OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: TARANAKI: Mt Egmont, Stratford Mountain House, P.R. Johnston, 12 Dec 1989 (PDD 56745). TAUPO: Tongariro National Park, Ketetahi Track, J. Bedford, 25 Jan 1985 (PDD 47823, OSC). REMARKS: Owing to the unique sporocarp structure and small spores, we prefer to place this taxon in a genus unto itself as Cunningham originally proposed. Apparently saprophytic on decaying wood. PROTUBERA MOLLER, IN SCHIMPER, BOT. MITTHEIL. AUS DEN TROPEN 7: 10, Basidiomata globose to depressed. Peridium thick and gelatinised, not easily separable from gleba, white, pale pink to pale red, surface glabrous to finely tomentose, occasionally cerebriform. Gleba pale grey olive to dark greyish yellow brown, with elongate empty locules. Odonr nil to sharply putrid. Basidia in a euhymenium, hyaline, thin-walled, 6-8-spored. Spores ellipsoid, statismosporic, orthotropic, smooth, lacking an utricle. Sterigmal attachment inconspicuous to slight. In KOH hyaline to pale green singly. No distinctive reactions to Melzer's reagent. Development angiocarpic, hypogeous to epigeous. ETYMOLOGY: From Latin protuberare, to swell out, presumably referring to the protuberance-like form of the indehiscent sporocarp. TYPE SPECIES: Protubera maracuja M611er. REMARKS: Protubera is related to the Clathraceae by its small, smooth spores and sutures which divide the peridium into sections. The genus is widely distributed, having been recorded from North and South America, Africa, Asia, and Australia. The very gelatinous nature ofp. parvispora and P. hautuensis makes peridial details of these species difficult to discern in KOH mounts of heat-dried specimens. Key to Protubera species 1 Epicutis separable from subcutis, spores ~m; hypogeous under Nothofagus P. nothofagi Epicutis not separable from subcutis, spores x ~tm; often epigeous or buried in rotting plant matter Gelatinised subcutis 5 mm thick, sutures conspicuous and common... P. parvispora Gelatinised subcutis <1 mm thick, sutures inconspicuous and occasional... P. hautuensis Protubera hautuensis Castellano & Beever, sp. nov. = Hysterangium hautu Cunningham, Trans. Roy. Soc. New Zeal. 67: 409, nom. nud. (Fig. 10) Basidiomata usque ad 4 cm diam., globosa vel depressa, alba vel lactea, contundentibus pallide rosea vel pallide lateritia, minute tomentosa. Gleba gelatinosa, pallide olivaceo-cinerea vel cinereoolivacea, loculis elongatis, vacuis. Rhizomorpha singularis, usque ad 2 mm diam., peridio concolor. Columella dendroidea, gelatinosa, translucens, truncata. Peridium fragile, non facile separabile, ram crassum; epicute ram crassa, hyphis hyalinis, intertextis, inflatis, 5-60 ~n diam.; subcute ram crassa, hyphis hyalinis intertextis, 24 ram diam., in matrice gelatinosa. Cutis suturis infrequentibus, hyphis hyalinis, intertextis, 2-5 ram diam. Trama usque ad 80 ram crassa, hyphis hyalinis, intertextis, 2-3 ram, in matrice gelatinosa. Sporae laeves, x ram, ellipsoideae, apice obtuso; paries minus quam 0.5 ram crassus; singulariter hyalinae vel pallide virides, aggregate pallide virides. Holotypus PDD Basidiomata up to 4 cm diam., globose to depressed, slightly ridged, white to bluish white when fresh, bruising pale pink to pale grey red, especially on ridges, surface finely tomentose, tomentum easily rubbing off upon handling; EtOH red to reddish brown. Gleba gelatinous, pale olive grey to greyish olive; locules elongate, empty. Rhizomorph stout, single, soon branching, up to 2 mm diam., concolorous with peridium, attached at base. Colume la distinct, dendroid, gelatinous, translucent. Taste not noted. Odour not noted. Peridinm fragile when dried, not easily separable from gleba, ~trn thick, 2-layered; epicutis

20 Castellano & Beever--Truffle-like fungi 323 Fig. 10 Protubera hautuensis. A, Basidiomata exterior and in section (PDD 52010). B, Detail of basidioma in section (PDD 52010). C, Basidiospores by bright field (PDD 52010). D, Basidiospores by SEM (PDD 52010). Scale: 1 cm (A), 1 mm (B), 10 Iam (C), 1 pm (D) pm thick, of hyaline, thin-walled, interwoven, inflated hyphae, 5-60 ~Jm diam., clamp connections absent; subcutis lain thick, of hyaline, interwoven hyphae, 2-4 pm diam., in a gelatinised matrix, clamp connections absent. Sutures infrequent, of hyaline, interwoven hyphae, 2-5 Iam diam., without clamp connections, connecting some locules to epicutis. Trama up to 80 ~Jm thick, of hyaline, thin-walled, interwoven hyphae, 2-3 ~rn diam., in a gelatinised matrix, clamp connections absent. Basidia not observed, "8 spored" (Cunningham). Spores smooth, x ~arn, ellipsoid; apex obtuse, base obtuse; wall <0.5 pm thick. Utricle absent. Spore eolour in KOH hyaline to pale green singly, pale green in mass. Habitat epigeous and buried amongst decaying leaves and white-rotted branches in podocarpbroadleaf and Nothofagus forests. Season February through October. Distribution New Zealand. ETYMOLOGY: From Maori Hautu, the district where the type was collected. TYPE: NEW ZEALAND, TAUPO, Waimarino,

21 324 New Zealand Journal of Botany, 1994, Vol. 32 J.C. Neill, Feb 1930 (holotype PDD 8311; isotypes DAOM , K, OSC) (type of Hysterangiunz hautu). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: AUCKLAND: Waitakere Ranges, Glen Esk Valley, B.E.G. Molesworth, 19 Aug 1947 (PDD 5581, OSC); Waitakere Ranges, Kauri Grove Track, R.P. Korf & R.E. Beever 83, 13 Aug 1981 (PDD 52007); same locality, R.E. Beever 85, 12 Sep 1981 (PDD 52009); same locality, R.E. Beever 598, 27 May 1986 (PDD 52010, OSC). COROMANDEL: Thames, Kauaeranga Valley, S.D. Baker, 26 Aug 1954 (PDD 13629, OSC); Coromandel Forest Park, track to Tarawaere Dam, L.E. Hooper, 1 Sep 1986 (PDD 51929). TAUPO: Urewera, Te Awhiti,?collector, Jul 1948 (PDD 6229, OSC); Turangi, Pihanga, J.M. Dingley, 10 Oct 1968 (PDD 45877, OSC); Karioi, Rotokuru Ecological Reserve, D.C. Hogan, 7 Sep 1987 (PDD 55124). REMARKS: Cunningham proposed the name Itysterangium hautu without a Latin description after 1935, so it is a nomen nudum in accordance with Article 36.1 of the Botanical Code. We validate the specific epithet, as hautuensis, in accordance with the code recommendations on forming epithets from place names. The occurrence of sutures through the gelatinised subcutis places this species in genus Protubera. The combination of thin peridial layers, infrequent sutures, and peridium colouring, especially upon handling, separates P. hautuensis from all other Protubera species. The record ofh. hautu from Queensland by Cribb (1958) is erroneous and based on a distinct, as yet unnamed, taxon. Protubera nothofagi Castellano & Beever sp. nov. (Fig. 11) Basidiomata usque ad 3 cm diam., globosa, depresse globosa vel irregularia, alba vel pallide luteorosea, contundentibus pallide avellanea, glabra. Gleba gelatinosa, obscure cinereo-olivacea vel obscure cinereo-luteobrunnea, loculis elongatis, plus minusve radiantibus, partialiter repletis. Rhizomorpha evanescens, singularis, crassa, peridio concolor. Columella dendroidea, gelatinosa, translucens. Sapor destitutus vel parum piperatus. Odor destitutus acute putridus. Peridium separabile, plus minusve 1 mm crassa; epicute facile e subcute separanti, (-350) pm crassa, hyphis hyalinis, irregularibus, intertextis vel subparallelis, 5-8 ]am diam., interdum usque ad 12.5(-25) pm inflatis; subcute plus minusve 800 pm crassa, hyphis hyalinis, irregularibus, intertextis, plus minusve 5 pm diam., in matrice gelatinosa. Cutis suturis frequentibus, epicutem interiorem simulantibus. Trama hyphis hyalinis, intertextis, plus minusve 5 pm diam., in matrice gelatinosa. Basidia x 4-9 ~m. Sporae laeves, 6-7 x pm, ellipsoideae, apice obtuso; paries minus quam 0.5 pm crassus; singulariter pallide virides, aggregate pallide olivaceo-virides. Holotypus PDD Basidiomata up to 3 cm diam., globose to depressed, larger sporocarps somewhat irregular, surface white to pale yellow pink when fresh, pale pink to brown when bruised, pale greyish yellow brown to pale yellow brown when dried, glabrous. Gleba gelatinous, dark greyish olive to yellow brown to olive black when deliquescent; locules elongate, more or less radially arranged, partially filled. Rbizomorph evanescent, single, stout, concolorous with peridium. Columella dendroid, up to 6 primary branches in longitudinal section, gelatinous, translucent to white near attachment point, translucent brown when dried. Taste nil to slightly peppery. Odour nil to sharply putrid when gleba deliquescent. Peridium 2-layered, c. 1 mm thick, epicutis readily peeling from subcutis; epicutis (-350) pm thick, of thin-walled, compact, irregularly shaped, subpericlinal to interwoven hyphae, 5-8 pm diam., occasionally inflated to 12.5(-25) pm diam., outer pm golden brown, inner portion hyaline, clamp connections absent; subcutis c. 800 ~am thick, of hyaline, irregularly shaped, compact, interwoven hyphae, c. 5 ~m diam., in a gelatinised matrix, clamp connections absent. Sutures transecting subcutis, common, structurally similar to, and continuous with, the inner portion of the epicutis and trama. Tralna of hyaline, thin-walled, compact, interwoven hyphae, c. 5 pm diam., in a gelatinised matrix of different texture to that of peridial subcutis, clamp connections absent. Basidia hyaline, thinwalled, elongate to mostly napiform, x 4-9 pm, broader at base than apex, 6-8-spored. Spores smooth, 6-7 x pm, ellipsoid, apex obtuse, base with a short sterigmal attachment to sessile; wall <0.5 pm thick. Utriele absent. Spore eolour in KOH pale green singly, pale olive green in mass. Habitat hypogeous in Nothofagus forest, putatively mycorrhizal with Nothofagus fusca, N. menziesii, N. solandri, and N. solandri var. cliffortioides. Season December through May. Distribution New Zealand. ETYMOLOGY: Referring to the association of this fungus with Nothofagus. TYPE: NEW ZEALAND, NELSON, Nelson City, track to Fringed Hill, R.E. Beever 667, 30 Apr 1987 (holotype PDD 48806; isotype OSC).

22 Castellano & Beever--Truffie-like fungi 325 Fig. 11 Protubera nothofagi. A, Basidiomata exterior and in section (PDD 48806). B, Detail of peridial wall in section (PDD 48806). C, Basidiospores by bright field (PDD 48806). D, Basidiospores by SEM (PDD 48806). Scale: 1 cm (A), 1 mm (B), 10 gm (C, D), OTHER COLLECTIONS EXAMINED: NEW ZEALAND: TAUPO: Turangi, M. Chu-Chou, 22 Apr 1981 (NZFR13296b); Kaimanawa Forest Park, Kiko Road, R.E. Beever 708, 7 May 1987 (PDD 48805, OSC); Kaimanawa Forest Park, Tree Trunk Gorge, R.E. Beever877, 19 May 1989 (PDD 55919); Tongariro National Park, Taranaki Falls Track, R.E. Beever 881, 20 May 1989 (PDD 55922). NELSON: Nelson City, track to Fringed Hill, J.D. Hayward & R.E. Beever 577, 28 Apr 1986 (PDD 48807, OSC); Takaka River, Cobb Dam road, Asbestos Forest Walk, M.K. Thayer, 2 Jan 1985 (PDD 48808, OSC). BULLER: Maruia, R.F.R. McNabb, 24 Apr 1969 (PDD 31435, OSC); Warbeck Scenic Reserve, Maruia Saddle, R.E. Beever 209, 16 Apr 1983 (PDD 48801, OSC); south of Lewis Pass, M.K. Thayer, 24 Dec 1984 (PDD 48809, OSC); NORTH CANTERBURY: Arthur's Pass National Park, Halpin Creek, Y. Doi & R.E. Beever 632, 22 Apr 1987 (PDD 48804). FIORDLAND: Fiordland

23 326 New Zealand Journal of Botany, 1994, Vol. 32 National Park, Kepler Track, R.E. Beever 999, 17 May 1990 (PDD 57835). REMARKS: Protubera nothofagi differs from other Protubera species in its larger spores, separable epicutis, and strict association with Nothofagus spp. Protubera parvispora Castellano & Beever sp. nov. (Fig. 12) Basidiomata usque ad 5.2 cm diam., globosa vel depressa, alba, contundentibus brunnea, glabra, cerebfiformia. Gleba gelatinosa, olivaceo-brunnea vel cinereo-olivacea, loculis usque ad 1 mm diam., elongatis, vacuis. Rhizomorpha singularis, usque ad 2 mm diam., peridio concolor. Columella dendroidea, plus minusve 1 mm diam., gelatinosa, translucens vel caesiella. Peridium non facile separabile, 5-6(-10) mm crassum; epicute lain crassa, hyphis hyalinis intertextis, irregulafibus, 5-9 lam diam., prope paginam 2-3 tam diam. et hyphis frequentibus, erectis, contractis e pagina projectis; subcute 5~5(-10) mm crassa, hyphis hyalinis irregularibus laxe intertextis, 1-3 0m diam., in matrice gelatinosa; fibulae communes, cutis suturis frequentibus, hyphis hyalinis, brunneolis vel brunneis, laxe intertextis, 1-3 lam diam. transecta. Trama hyphis hyalinis, laxe intertextis, 1-3/am diam., in matrice gelatinosa. Sporae laeves 4(-5) x 1.5(-2) 0m, ellipsoideae, apice obtuso; paries minus quam 0.5 pm crassus; singulafita pallide virides, aggregatae virides. Holotypus PDD Basidiomata up to 5.2 cm diam., globose to depressed, white when fresh, bruising brown, white to pale yellow brown when dried, surface glabrous and with a cerebriform appearance; KOH and EtOH negative. Gleba gelatinous, olive brown to greyish olive; locules up to l mm diam., elongate, empty. Rhizomorph single but soon branching, up to 2 mm diam., concolorous with peridium. Columella dendroid, c. 1 mm diam., gelatinous, translucent to grey blue. Taste not noted. Odour nil to faint. Peridium not easily separable from gleba, 5-6 (-10) mm thick, 2-layered; epicutis pm thick, of hyaline, thin-walled, compact, irregularly shaped, interwoven hyphae, 5-9 pm diam., much finer near peridial surface, 2-3 ~m diam., with frequent, erect, tapered hyphae projecting from surface, clamp connections absent; subcutis 5-6 (-10) mm thick, of hyaline, loosely interwoven, irregularly shaped hyphae, 1-3 lain diam., in a gelatinised matrix, clamp connections common. Sutures transecting subcutis, frequent, of mostly hyaline, occasionally pale brown to brown, loosely interwoven hyphae, 1-3 om diam., contiguous with epicutis, and extending intermittently along interface between gleba and subcutis. Trama of hyaline, thin-walled, loosely interwoven to subparallel hyphae, 1-3 gm diam., in a gelatinised matrix, clamp connections absent. Basidia not observed. Spores smooth, 4(-5) x 1.5(-2) pro, ellipsoid, apex obtuse, base with slight sterigmal attachment to sessile; wall <0.5 om thick. Utriele absent. Spore eolour in KOH pale green singly, green in mass. Cultures grown from glebal tissue transferred to malt extract agar at 20~ produced slow-growing (1.6 mm radial growth/week) colonies; after 17 weeks, cultures showed an irregular periphery, with submerged, hyaline hyphae marginally and felted, white aerial hyphae centrally. Habitat epigeous on, to slightly immersed in, rotting wood and other plant material, sometimes on soil, in podocarp-broadleaf forest. Season April through November. Distribution New Zealand. ETYMOLOGY: From Latin parvus, small, and sporum, spore, referring to the small spores. TYPE: NEW ZEALAND, COROMANDEL, Coromandel Forest Park, track to Kaitarakihi, R.E. Beever 465b, 13 May 1985 (holotype PDD 52011, isotype OSC). OTHER COLLECTIONS EXAMINED: NEW ZEALAND: AUCKLAND: Hunua Range, Moumoukai Valley, J.M. Dingley, 14 Jul 1946 (PDD 4579, OSC); Hunua Range, Mangatawhiri, J.M. Dingley, 2 Oct 1946 (PDD 5111, OSC). WAIKATO: Mt Te Aroha, G.H. Cunningham, 9 Sep 1954 (PDD 13628, OSC). BAY OF PLENTY: Rotorua, Lake Okataina, S. Rowe, 1 Nov 1988 (PDD 56744). TAUPO: Urewera, Te Awhiti,?collector, Jul 1948 (PDD 6228, OSC); Tongariro National Park, Mangawhero Forest Walk, G.L. Barron, 20 May 1989 (PDD 55932, OSC); Tongariro National Park, vicinity Ohakune ranger station, L. Gibbons, 20 May 1989 (PDD 55930, OSC). REMARKS: Protubera parvispora has smaller spores than the Australian P. canescens Beaton & Malajczuk and South African P. africana Lloyd, with which it is macroscopically aligned. Two collections [AUCKLAND: Hunua Range, Moumoukai Valley, J.M. Dingley, 11 Jun 1949 (PDD 7651, OSC). SOUTHLAND: Maclennan, J.R.J. Moore, Apr 1937 (PDD 8312, OSC)] generally match this species but differ somewhat in peridial characters. UNCERTAIN TAXA 1. Phallogaster globosus Lloyd, Mycol. Writings 5: 739, TYPE: NEW ZEALAND, DUNEDIN, Caversham, W. Scarfe (Lloyd 22951),?date (holotype BPI).

24 Castellano & Beever--Truffle-like fungi 327 Fig. 12 Protubera parvispora. A, Basidiomata exterior and in section (PDD 52011). B, Detail of attachment region in section (PDD 52011). C, Basidiospores by bright field (PDD 52011). D, Basidiospores by SEM (PDD 52011). Scale: 1 cm (A), 1 mm (B), 10 }am(c), 1 lam (D).

25 328 New Zealand Journal of Botany, 1994, Vol. 32 REMARKS: This taxon was described from one basidioma forwarded to Lloyd in formalin. The specimen has since been dried, and microscopic details have proved difficult to resolve. Lloyd's description and illustration indicate that in general appearance it resembles Protubera hautuensis and P. parvispora. It appears, however, to differ from both in microstructure, the salient features of which we determined as follows: peridium 2-layered, epicutis thin, hyphae 3-4 ram diam. protruding from surface, clamp connections present, pale brown, interwoven underlying layer intergrading into subcutis of inflated cells up to 35 x 20 ~tm, sutures present; glebal trama of compact, interwoven hyphae 2-5 ~tm diam., larger near subcutis, mostly collapsed, some capitate to 12 lam diam. at apex; spores 4-5 x lam. It does not fit comfortably in Phallogaster, which otherwise comprises one Northern Hemisphere species, Phallogaster saccatus Morgan, characterised by a distinct, sterile, stipitate base and flaking peridium. We suspect that it represents a species of Protubera, but in the absence of further collections and information on its fresh appearance we make no decision. We disagree with Cunningham (1944), who suggested that the species was based on a phalloid egg "probably that of Clathrus cibarius". 2. Hysterangium sp 1 and sp 6 of Chu-Chou & Grace (1983) are considered not to belong in Hysterangium or any other genus covered in this paper. ACKNOWLEDGMENTS To Jim Trappe for encouragement, advice, and providing the Latin diagnoses; to Kylie Head for SEM and help in preparing the plates; the photographic section of Mt Albert Research Centre for photographic assistance; curators of herbaria for loan of collections; and many friends and colleagues who assisted with specimen collection and field logistics. M.A.C. acknowledges support from National Science Foundation Grant BSR R.E.B. acknowledges support from Foundation for Research, Science and Technology contract CO9309, and the New Zealand - United States Co-operative Science Programme. REFERENCES Beaton, G.; Pegler, D. N.; Young, T. W. K. 1985a: Gasteroid basidiomycota of Victoria State, Australia: 8-9. Kew bulletin 40: Beaton, G.; Pegler, D. N.; Young, T. W. K. 1985b: Gasteroid basidiomycota of Victoria State, Australia: 4. Hysterangium. Kew bulletin 40: Chu-Chou, M.; Grace, L. J. 1983: Hypogeous fungi associated with some forest trees in New Zealand. New Zealand journal of botany 21: Cribb, J. W. 1958: The gasteromycetes of Queensland - IV. Gautieria, Hysterangium and Gymnoglossum. University of Queensland papers, Department of Botany 3: Crosby, T. K.; Dugdale, J. S.; Watt, J. C. 1976: Recording specimen localities in New Zealand: an arbitrary system of areas and codes defined. New Zealand journal of zoology 3:69 + map. Cunningham, G. H. 1924: The development of Gallacea scleroderma (Cke.) Lloyd. Transactions of the British Mycological Society 9: Cunningham, G. H. 1934: The gasteromycetes of Australasia. XVI. Hymenogastraceae, Part I: the genera Rhizopogon, Melanogaster and Hymenogaster. Proceedings of the Linnean Society of New South Wales 59: Cunningham, G. H. 1944: The gasteromycetes of Australia and New Zealand. Dunedin, John McIndoe. 236 p. Holmgren, P. K.; Holmgren, N. H.; Barnett, L. C. 1990: Index Herbariorum. Part 1: The herbaria of the world. 8th ed. Regnum Vegetabile Vol Horak, E. 1973: Fungi agaricini novaezelandiae II. Thaxterogaster. Beihefie zur nova hedwigia 43: Jtilich, W. 1981: Higher taxa of basidiomycetes. Bibliotheca mycologica 85: Patouillard, N. 1910: Champignons de la Nouvelle- Caledonie VI. Le genre Gallacea Lloyd. Bulletin de la Societe Mycologique de France 27: Singer, R.; Smith, A. H. 1958: Studies on secotiaceous fungi III. The genus Weraroa. Bulletin of the Torrey Botanical Club 85: Zeller, S. M.; Dodge C. W. 1929: Hysterangium in North America. Annals of the Missouri Botanical Garden 16: