MycoKeys 7: 3 1-44 (20 | 3) A peer-reviewed open-access journal doi: 10.3897/mycokeys.7.4710 RESEARCH ARTICLE O Myco Keys www.pensoft.net/journals/mycokeys Launched to accelerate biodiversity research Three new species of foetid Gymnopus in New Zealand Jerry A. Cooper', Pat L. Leonard? | Landcare Research, PO Box 40, Lincoln, 7640, New Zealand 2. Motueka, New Zealand Corresponding author: Jerry Cooper (cooperj@landcareresearch.co.nz) Academic editor: S. Redhead | Received 18 January 2013 | Accepted 24 June 2013 | Published 26 June 2013 Citation: Cooper JA, Leonard PL (2013) Three new species of foetid Gymnopus in New Zealand. MycoKeys 7: 31-44. doi: 10.3897/mycokeys.7.4710 Abstract We describe three new species, Gymnopus imbricatus, G. ceraceicola and G. hakaroa, from New Zealand that are similar to G. foetidus (= Micromphale foetidum), growing on wood, with an insititious stipe and foetid odour. The position of these species within the /gymnopoid clade is confirmed by ITS sequence analysis. Key words Gymnopus, Micromphale, New Zealand Introduction The new species we describe are members of the family Omphalotaceae Matheney et al. (2006) and have the morphological characteristics of Gymnopus section Véestipedes subsection /mpudicae (Antonin & Noordeloos 2010) which contains Micromphale foetidum (Sowerby) Singer, the type species of the formerly recognised genus Microm- phale (e.g. in the sense of Singer 1986). Fruitbodies of the group often have a foetid odour when crushed, described as like rotting cabbage or garlic. The only existing re- cords of this group in New Zealand were found to be misapplications of names applied to northern hemisphere species. Moncalvo et al. (2002) investigated nLSU rDNA sequence data for a large number of agarics and recognised a /micromphale clade containing M. foetidum (AF261328). The clade also contained Gymnopus pro parte, Caripia, Setulipes and Micromphale. Copyright J.A. Coope, PL Leonard. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 32 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) Their /micromphale clade was nested within a broader /lentinuloid clade including Rhodocollybia, Marasmiellus ramealis (Bull.) Singer, Marasmius scorodonius (Fr.) Fr. and Lentinula. Mata et al. (2004), based on an LSU analysis, also identified a clade containing sequences of M. foetidum and material named as Setulipes androsaceus (L.) Antonin and Gymnopus fusipes (Bull.) Gray, the type species of those respective gen- era. They adopted a broad concept of Gymnopus incorporating these genera together with Marasmiellus. Wilson and Desjardin (2005) used LSU to examine the group and identified a /gymnopus clade containing G. fusipes, M. foetidum, S. androsaceus at its core with Micromphale perforans (Hoftm.) Gray lying on its boundary. These results were broadly supported by Mata et al. (2006) in their analysis using [TS1—5.8-ITS2 but they demonstrate clustering of G. fusipes, S. androsaceus and Micromphale on the periphery of a concentration of Gymnopus-labelled samples. On the basis of these re- sults the currently generally accepted concept of Gymnopus is broad (e.g. Noordeloos 2012), and incorporates a number of previously recognised genera. Hughes et al. (2010) erected the genus Connopus to accommodate the Gymnopus acervatus group within the gymnopoid clade and presented LSU and ITS data indicating its place- ment close to RKhodocollybia. Their LSU analysis supports a core gymnopoid clade containing G. fusipes, S. androsaceus, which once again places Micromphale foetidum and M. perforans on a boundary with a sister group containing Rhodocollybia, Mar- asmiellus juniperinus Murrill and various Gymnopus species. The /gymnopus, clade as interpreted by Hughes et al., contains significant substructure. A multi-gene analysis including more representatives may indicate the recognition of further segregates at genus-level. In this paper we accept our newly described species within the current broad concept of Gymnopus whilst recognising their close alliance to the historical concept of the genus Micromphale. For this study we analysed ITS1—5.8-ITS2 data for related New Zealand collec- tions together with representative sequences from Genbank, many from the studies cited above. The structure of our ITS tree is consistent with these previous analyses, and once again identifies a /micromphale clade closely linked to core Gymnopus spe- cies. ITS data generated for a number of representative collections of our newly de- scribed taxa support species concepts based on morphology. Materials and methods Morphological protocols Spore dimensions are stated as the mean + 1.5 SD of 20 measurements, thus covering 86% of measurements under an assumed normal distribution model. Fresh or dried material was examined mounted in 10% KOH or Melzer’s reagent. Material was hand- sectioned. Some micrographs were obtained under DIC conditions. Measurements were always taken without DIC optics and an extended objective iris in order to max- imise boundary contrast. Three new species of foetid Gymnopus in New Zealand 33 Phylogenetic protocols DNA extraction and sequencing followed the protocols outlined in Cooper and Leon- ard (2012). We downloaded from Genbank selected sequences used in cited publica- tions, together with close BLAST matches, Table 1. General sequence management was carried out using Geneious (Drummond et al. 2011). Data exchange between ap- plications was facilitated using Alter (Glez-Pefa et al. 2010). Sequence alignment was carried out using MAFFT within Geneious (Katoh et al. 2002). A maximum likeli- hood analysis was executed using RAxML (Stamatakis 2006), with 100 bootstrap runs, launched from Topali 2.5 (Milne et al. 2004). The substitution model of GTR+G was recommended by Topali 2.5. We selected a sequence of Anthracophyllum archeri (Berk.) Pegler as the outgroup. Table |. ITS Sequences used in the analysis. New sequences generated for this analysis are in bold. Genbank # PDD Voucher# | Country DQ444308 TENN50049 — | Anthracophyllum archeri New Zealand DQ480112 TENN58672 Gymnopus alkalivirens Greenland DQ480114 TENN55834 Gymnopus alpinus Scotland AY256691___ | TENN57012_|Gymnopusaguous |_| Germany DQ449971_ | TENN59738_| Gymnopusaquosus |S KC248409 PL6304 Gymnopus ceraceicola PDD 101750 New Zealand KC248389 PL126406 Gymnopus ceraceicola PDD 101754 New Zealand KC248400 PL189402 Gymnopus ceraceicola PDD 76358 New Zealand KC248403 PDD 80771 New Zealand KC248405 Gymnopus ceraceicola PDD 87181 New Zealand KC248404 Gymnopus ceraceicola PDD 87424 New Zealand KC248394 Gymnopus ceraceicola PDD 87483 New Zealand KC248408 Gymnopus ceraceicola PDD 87661 New Zealand KC248392 PDD 90101 New Zealand KC248391 KWH12891 Gymnopus ceraceicola PDD 90119 New Zealand KC248393 RHP12871 Gymnopus ceraceicola PDD 90132 New Zealand KC248397 JAC11005 Gymnopus ceraceicola PDD 95459 New Zealand KC248395 JAC11093 PDD 95544 New Zealand AY256690 TENN57012 Gymnopus dryophilus _—— USA DQ449974 TENN58087 Gymnopus dryophilus Costa Rica AF505778 TENN 59141 — | Gymnopus dysodes Costa Rica AY256694 TENN59457 | Gymnopus earleae USA DQ449973 TFB10718 Gymnopus exculptus [ee] Greenland AF505780 FB11434 Gymnopus foetidum oe, USA AY256710 TENN59217 Gymnopus fusipes France KC248407 JAC9585 Gymnopus hakaroa PDD 81086 New Zealand KC248410 JAC10225 Gymnopus hakaroa PDD 87315 New Zealand KC248411 PL25404 PDD 101753 _| New Zealand KC248406 JAC10089 Gymnopus imbricatus PDD 87186 New Zealand 34 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) Genbank # Country KC248401 JAC10322 Gymnopus imbricatus PDD 87410 New Zealand KC248399 PDD 87659 New Zealand KC2ABkO2 PDD 87660 [New Zealand KC248396 JAC10495 Gymnopus imbricatus PDD 87675 New Zealand KC248390 JAC11038 Gymnopus imbricatus PDD 95489 New Zealand AS05779 TENN56658 Gymnopus impudicus Costa Rica DQ449986 | Duke RV94154 | Gymnopus iocephalus | _LUSA AY256693___|TENN59532_| Gymnopusjunguilleus_ |__| USA DQ449960 TENN50620 Gymnopus ocior Switzerland DQ449972 TENN56321 Gymnopus subsulphureus USA AY263453 AWW115 Gymnopus vitellinipes Java/Bali AY256708___|TENN59540__|Marasmiellus juniperinus_ |__| USA GU234007 JB14 Marasmius androsaceus Sweden DQ444312 TENN50482 — | Marasmius androsaceus UK DQ444311 TENN50704 — | Marasmius androsaceus USA DQ449990__ | TENN59293__| Micromphale brassicolens_ |__| Austria Results Our analysis places the New Zealand taxa in a monophyletic clade close to G. foetidum and G. brassicolens historically recognised in the genus Micromphale (Fig. 1). The com- bination of sequence data and morphological analysis of many collections indicate two major groups which we equate with the newly described species G. imbricatus and G. ceraceicola. In addition we recognise a further species, G. hakaroa, which is poorly dis- tinguished from G. imbricatus on the basis of ITS sequences but which is morphologi- cally consistently different. Minor sequence variation in the G. ceraceicola group does not correlate with morphology and we choose to recognise these specimens as a single species. More information and images of collections may be found on the Landcare Research website (Systematics Collections Data). Gymnopus ceraceicola J.A. Cooper & P. Leonard, sp. nov. http://species-id.net/wiki/Gymnopus_ceraceicola Holotype: PDD 87181. Registration identifier: IF550091 Diagnosis. G. ceraceicola is distinguished from related New Zealand species by the combination of pruinose, central stipe and dark pileus. Macromorphology. Pileus 5—20 mm, generally broadly convex to applanate, but sometimes campanulate when young, brick to purplish chestnut, minutely felty, ra- dially furrowed and striate towards the margin, margin slightly fimbriate. Lamellae cream, creamy yellow to vinaceous buff, waxy, adnate. Lamellae present, in series of Three new species of foetid Gymnopus in New Zealand a2 Anthracophyllum archeri/TENN50049 Marasmius androsaceus|TENN50704 Marasmius androsaceus|JB14 Marasmius androsaceus|TENN50482 Gymnopus fusipes|TENN59217 Marasmiellus juniperinus|TENN59540 Gymnopus vitellinipes|AWW115 Gymnopus alkalivirens|TENN58672 Gymnopus earleae|TENN59457 Gymnopus dryophilus|TENN57012 Gymnopus ocior/TENN50620 Gymnopus aquosus|TENN57012 Gymnopus subsulphureus|TENN56321 Gymnopus alpinus|TENN55834 Gymnopus dryophilus|TENN58087 Gymnopus exculptus|TFB10718 57 Gymnopus junquilleus|TENN59532 Gymnopus aquosus|TENN59738 Gymnopus impudicus|TENN 56658 Gymnopus iocephalus|Duke RV94154 98 Gymnopus dysodes|TENN 59141 Micromphale brassicolens|TENN59293 Micromphale foetidum|FB11434 Gymnopus imbricatus|JAC10816 88 Gymnopus imbricatus|JAC11038 Gymnopus imbricatus|}JAC10322 Gymnopus imbricatus|JAC10495 69 Gymnopus imbricatus|JAC10310 Gymnopus imbricatus|JAC10089 Gymnopus imbricatus|pl25404 Gymnopus imbricatus|JAC10815 Gymnopus hakaroa|JAC9585 i 98 Gymnopus hakaroaWAC10225 Gymnopus ceraceicolajJAC1 1093 Gymnopus ceraceicola|RHP12871 Gymnopus ceraceicola|KWH12891 Gymnopus ceraceicola|PL126406 Gymnopus ceraceicolajJAC11005 Gymnopus ceraceicola|pl6304 Gymnopus ceraceicola)JAC10084 x Gymnopus ceraceicola|PL189402 20 differences Gymnopus ceraceicolajJAC9334 — Gymnopus ceraceicolajWAC10336 Gymnopus ceraceicolajJAC10817 Gymnopus ceraceicolajJAC10395 Gymnopus ceraceicola|RHP13063 Figure |. Maximum likelihood cladogram of selected ITS sequences, with bootstrap proportion. Red bar = Gymnopus imbricatus, green bar = G. hakaroa, blue bar = G. ceraceicola three: intercalated short/long/short. Stipe central, cartilaginous, 10-20 x 1-2 mm, equal, brown vinaceous, sometimes paler towards apex or base, always entirely finely pruinose. Stipe base insititious and always associated with a thin waxy to chalky cream layer of partially gelatinised hyphae covering the substrate. This layer is often extensive, with a distinct margin, and often green with algal cells. Fruitbodies with garlic/rotten cabbage smell, especially when crushed. Micromorphology. Pileipellis a partially gelatinised radially arranged clamped cutis of smooth hyphae to 5 um diameter, with brown extra-cellular encrustation. Epi- dermal layer to 140 um. Subepidermis of thick glassy—walled non-gelatinised smooth hyaline hyphae, weakly dextrinoid. Basidia clavate to 40 x 8 um. Sterigmata to 7 um, 4—spored. Basidioles cylindrical, tapering towards apex, 40 x 4 um. Spores hyaline, lacrymoid, 7.9 + 1 x 4.5 + 0.6 um, Q = 1.8 + 0.1 including apiculus. Cheilocystidia and pleurocystidia not observed. Stipitipellis a cutis of brown parallel hyphae, to 5 um wide. Caulocystidia smooth, hyaline, agglutinated into fascicles. 36 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) PY ies: Tek. - es. h BE ae Figure 2. Gymnopus ceraceicola Holotype, PDD 87181. Fruitbodies. Figure 3. Gymnopus ceraceicola Holotype, PDD 87181. A Spores (KOH) B Agglutinated fascicles of caulocystidia on stipe (KOH). Habitat. Colonies of a few to hundreds of fruitbodies on bark of fallen, dead branches and twigs, especially Nothofagus. Distribution. Broadly distributed and common in both North and South Islands of New Zealand. Etymology. Ceraceicola, indicating association with a basal waxy layer, although this feature is common to the three species described here. Notes. Sequence data indicate variability in the taxon but the morphological de- tails are constant and we choose to recognise a single species. New Zealand records of Gymnopus (Micromphale) foetidum and Gymnopus (Micromphale) brassicolens are at- tributable to G. ceraceicola. Authentic New Zealand material of these two species has not been identified. Gymnopus brassicolens has paler pileus colours, non-gelatinized pileipellis, cheilocystidia and pileipellis elements with lateral projections, and larger basidiospores. Gymnopus foetidus is macroscopically similar but does not possess the agelutinate fascicles of caulocystidia of G. ceraceicola. Three new species of foetid Gymnopus in New Zealand af Specimens examined. New Zealand, North Island: PDD 40852, on dead wood, Anawhata Rd., Waitakare Ranges, Collector P.R. Johnston & G. Samuels, 9 June 1981. PDD 80771, on dead wood of Beilschmiedia tawa, Erua Forest, Taupo, Collector J.A. Cooper (JAC9334), 4 April 2005. PDD 87382, on dead wood of Nothofagus fusca, Mt Holdsworth, Gentle Annie Track, Wairarapa, Collector J.A. Cooper JAC10294), 11 May 2007. PDD 87483, on wood, Mt Holdsworth, Donnelly Flat Loop Track, Wairarapa, Collector G. Gates & D. Ratkowsky JAC10395), 7 May 2007. PDD 87424, on dead bark of Nothofagus, Mt Holdsworth, Gentle Annie Track, Wairarapa, Collector J.A. Cooper JAC10336), 11 May 2007. PDD 95544, on bark of Nothofagus fusca, Rimutaka Forest Park, Wellington, Collector J.A. Cooper JAC11093), 14 May 2009. PDD 95545, on bark of dead branch of Nothofagus fusca, Rimutaka Forest Park, Wellington, Collector J.A. Cooper JAC11094), 14 May 2009. New Zealand, South Island: PDD 76357, on dead twig of Nothofagus, Canaan Road Track, Nelson, Collector P.L. Leonard, 30 April 2002. PDD 96730, on dead wood, Wangapeka, Nelson, collector P.L. Leonard (PL126406), 14 April, 2006. PDD 90101=TENN 061068, on bark, vicinity of Seddonville, Charming Creek Track, Nel- son, Collector R.H. Petersen (RHP 13063), 11 May 2006. PDD 76358, on bark on dead branch of Nothofagus menziesii, Lake Daniels Track, Nelson, Collector P.L. Leon- ard (PL189402), 2 April 2002. PDD 95459, on bark of dead branch of Nothofagus so- landri, Kowai Bush, Springfield, Mid Canterbury, Collector J.A. Cooper JAC11005), 2 May 2009. PDD 95462, on bark of dead branch of Nothofagus solandri, Kowai Bush, Springfield, Mid Canterbury, Collector J.A. Cooper JAC11008), 2 May 2009. Holotype PDD 87181, on dead branch of Nothofagus fusca, Hinewai Reserve, Akaroa, Mid Canterbury, Collector J.A. Cooper (JAC10084), 3 June 2006. PDD 87661, on dead twigs of Leptospermum scoparium, Government Track, Waipori Falls Road, Dun- edin, Collector K. Soop (JAC10817), 12 May 2008. PDD 96636, on dead wood of Nothofagus solandri, Lake Hauroko, Fiordland, Collector P. White JAC12522). 7 May 2012. PDD 90119 =TENN061007, on twigs, Vicinity of Te Anau, Kepler Track from Rainbow Reach, Fiordland, Collector K.W. Hughes (KWH12891), 30 April 2006. PDD 90132=TENN060986, vicinity Manapouri, Borland Lodge, Nature Track, Fiordland, Collector R.H. Petersen (RHP 12871), 29 April 2006. Gymnopus imbricatus J.A. Cooper & P. Leonard, sp. nov. http://species-id.net/wiki/Gymnopus_imbricatus Holotype: PDD 95489. Registration identifier: IF550092 Diagnosis. G. imbricatus is distinguished from related New Zealand species by the smooth stipe, larger basidiospores, and imbricate habit. Macromorphology. Pileus 3-20 mm in diameter convex, cream to fawn, minute- ly felty, radially furrowed and striate towards the margin, margin fimbriate. Lamellae cream to creamy yellow, adnate. Lamellae present, in series of two: short/long. Stipe mostly eccentric, cartilaginous, to 3 x 0.5 mm, equal, umber to black, sometimes paler 38 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) , + ace ei ab hi at a Seis FT / eh ii | os } 7 yy = Figure 4. Gymnopus imbricatus. A Holotype PDD 95489. Fruitbodies, scale 1 cm B PDD 87186. Scale 1 mm. 20 um Figure 5. Gymnopus imbricatus Holotype PDD 95489. Spores (in KOH). towards base, always entirely smooth. Stipe base insititious and usually associated with a thin waxy to chalky cream layer of partially gelatinised hyphae covering the substrate, usually green with algal cells. Fruitbodies with garlic/rotten cabbage smell, especially when crushed. Micromorphology. Pileipellis a partially gelatinised irregular clamped cutis of hy- phae 4 um diameter, without intra or extracellular pigmentation, terminal layer with gelatinised coralloid elements, to 2 um wide, and occasional small finger-like tricho- dermal elements to 20 um. Epidermal layer to 25 um. Subepidermis of thick glassy- walled non-gelatinised smooth hyaline hyphae, weakly dextrinoid. Basidia clavate to 50 x 10 um. Sterigmata to 5 um, 4~spored. Basidioles to 50 x 6 um cylindrical and tapered towards apex. Spores hyaline, lacrymoid 9.8 + 1.2 x 5.1 + 0.4 um, Q= 1.9 + 0.3 including apiculus. Cheilocystidia and pleurocystidia not observed. Stipitipellis a cutis of parallel brown hyphae, to 6 um wide. Caulocystidia absent. Three new species of foetid Gymnopus in New Zealand ao Habitat. Forming imbricate colonies of dozens to hundreds of fruitbodies on bark and decorticate wood of dead branches and twigs, especially Kunzea and Leptospermum but occurs with other trees. Also occurs at the stem base of live trees. Distribution. Broadly distributed and common in both North and South Islands of New Zealand. Etymology. Imbricatus, pertaining to the often tiered and overlapping eccentri- cally stemmed caps. Specimens examined. New Zealand, North Island: PDD 80766, on bark of Beils- chmedia tawa, Erua Forest, Taupo, collector J.A. Cooper JAC9329), 4 April, 2005. PDD 87398, bark on dead branch of Nothofagus, Waiohine Gorge, Wairarapa, Collector J.A. Cooper JAC10310), 10 May 2007. PDD 87410, dead stems of Ripogonum scan- dens, Waiohine Gorge, Wairarapa, Collector J.A. Cooper JAC10322), 10 May 2007. New Zealand, South Island: PDD 101753, dead branches of Nothofagus menziesii, Riwaka Resurgence, Nelson, Collector P.L. Leonard (PL25404), 10 April, 2006. PDD 96141, dead twigs of Kunzea ericoides, Mt Fyffe Track, Kaikoura, collector J.A. Cooper (JAC11734), 26 Feb. 2011. PDD 80154, dead log of Nothofagus menziesii, Lewis Pass, Buller, collector J.A. Cooper JAC8287), 24 November, 2001. PDD 80157, on dead de-corticate log, Lyell Walkway, Nelson, collector J.A. Cooper (JAC80157), 25 No- vember, 2001. PDD 87675, living stem of Fuchsia excorticata, Saddle Hill, Mid Can- terbury, Collector J.A. Cooper JAC10495), 22 May 2005. Holotype PDD 95489 (Figs 4 and 5), base of live trees of Kunzea ericoides, Kennedy’s Bush, Mid Canterbury, Collector J.A. Cooper (JAC11038), 24 May 2009. PDD 79799, bark of dead tree, Kennedy’s Bush, Mid Canterbury, Collector J.A. Cooper JAC8921), 20 March 2004. PDD 87186, on bark of living tree of Kunzea ericoides, Hinewai Reserve, Akaroa, Mid Canterbury, Collector J.A. Cooper JAC10089), 3 June 2006. PDD 87660, fallen log, Racemans Track, Silverstream Valley, Dunedin, Collector S. Dodd (JAC10816), 13 May 2008. PDD 87659, on dead twigs of Kunzea ericoides, Evansdale Glen, Dunedin, Collector P.R. Johnston JAC10815), 12 May 2008. Gymnopus hakaroa J.A. Cooper & P. Leonard, sp. nov. http://species-id.net/wiki/Gymnopus_hakaroa Holotype: PDD 87315. Registration identifier: IF550093 Diagnosis. G. hakaroa is distinguished from G. ceraceicola by smaller stature and a pru- inose stipe lacking fascicles of agglutinate caulocystidia. It is distinguished from G. im- bricatus by non-imbricate growth, a consistently central stipe, and smaller basidiospores. Macromorphology. Pileus 3-10 mm diam. convex, rusty tawny to umber, mi- nutely felty, weakly radially furrowed and striate towards the margin. Lamella cream to yellow, waxy. Lamellae present, in series of three: intercalated short/long/short. Stipe central, cartilaginous, to 5 x 0.6 mm, equal, umber to black, paler towards base, smooth to minutely pruinose. Stipe base insititious and always associated with an obvi- ous waxy to chalky cream layer of partially gelatinised hyphae covering the substrate, 40 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) al q on sail At a . i > “ag 2 ! : ] ey ue 7 ; . J PA. \ ws : eo k . P . a 4 . & = a ‘ J 3 Vy 7 et | a Pe , a 4 7 me Boy ? Ponta sg ake ae Nae Figure 6. Gymnopus hakaroa A PDD 81086. Fruitbodies B scale= 2 mm Figure 7. Gymnopus hakaroa Holotype PDD 87315. Fruitbodies, showing waxy substratum. usually green with algal cells. Fruitbodies with garlic/rotten cabbage smell, especially when crushed. Micromorphology. Pileipellis a partially gelatinised radially arranged clamped cutis of smooth hyphae to 3 um in diameter, with brown extra-cellular encrustation. Epidermal layer to 80 um. Subepidermis of thick glassy-walled non-gelatinised smooth hyaline hyphae, to 3 um in diameter, weakly dextrinoid. Basidia clavate to 40 x 8 um. Three new species of foetid Gymnopus in New Zealand 4] Sterigmata to 7 um, 2—4-spored. Basidioles cylindrical and tapered towards apex 40 x 6 um. Spores hyaline, lacrymoid 8.3 + 1 x 4.8 + 0.3 um, including apiculus, Q = 1.7 + 0.2. Cheilocystidia and pleurocystidia not observed. Stipitipellis a cutis of hyaline to pale brown hyphae, to 5 um wide. Stipe without caulocystidia. Habitat. Forming imbricate colonies of dozens to hundreds of fruitbodies on de- corticate dead wood. Distribution. Currently G. /akaroa is only known from a single location on the Canterbury Port Hills in the South Island of New Zealand. Etymology. Hakaroa, a Maori name for the Bank’s Peninsula region of New Zealand. Notes. Sequence data (Fig 1) indicates a close phylogenetic relationship to G. im- bricatus but there are consistent and substantial morphological differences. Specimens examined. New Zealand, South Island: Holotype PDD 87315 (Figs 6 and 7) on dead log, Kennedys Bush Reserve, Port Hills, Mid Canterbury, Collec- tor J.A. Cooper JAC10225), 11 Feb. 2007. PDD 81086 (Fig. 8), on dead wood of Kunzea ericoides, Kennedys Bush Reserve, Port Hills, Mid Canterbury, Collector J.A. Cooper (JAC9585), 23 July, 2007. PDD 96390, on dead decorticate log of Melicy- tus ramiflorus, Kennedys Bush Reserve, Port Hills, Mid Canterbury, Collector J.A. Cooper (JAC11301), 17 April, 2010. Dicussion Gymnopus imbricatus, as its name suggests forms dense populations of small imbricate fruitbodies. It is most commonly associated with tea-tree (Kunzea ericoides and Lepto- spermum scoparium) and often found on the bark at the base of living trees. Gymnopus hakaroa is larger, with a dark minutely pruinose cap and again forms dense populations 42 Jerry A. Cooper & Pat L. Leonard / MycoKeys 7: 31-44 (2013) ie i eS MES Figure 9. Gymnopus hakaroa PDD 96390. Stipe base (arrow) with surrounding algal mat. Inset, primordial arising from algal mat. Figure 10. Gymnopus hakaroa PDD 96390. Pockets of algal cells embedded in hyphal tissue of stem base (cotton blue stain). Three new species of foetid Gymnopus in New Zealand 43 on the bark of dead logs. These two species have smooth stems. Gymnopus ceraceicola is distinguished by larger fruitbodies, a pruinose stipe, and is more commonly associated with southern-beech forests on dead fallen logs. The species of Gymnopus described here belong in the /micromphale clade of Moncalvo et al. (2002) and share the diagnos- tic feature of this clade of a foetid odour likely due to the presence of mercaptan-like compounds. In New Zealand this feature is shared with Mycetinis curraniae (G. Stev.) J.A. Cooper & P. Leonard, a marasmioid fungus distinguished by its ornamented hy- meniderm pileipellis. Another very distinctive character common to all three Gymnopus species, and visible in the accompanying photographs (Figs 2 and 6), is the presence of a waxy layer of partially gelatinised hyphae on the substrate from which the fruitbodies emerge. This layer is usually green from the presence of embedded algal cells. Interest- ingly, some published images of G. foetidus in the northern hemisphere also show a similar layer, e.g. Antonin and Noordeloos (2010). Detailed examination of our material does show algal cells deeply embedded within the context of the waxy layer and the basal portion of the stipe (Figs 9 and 10), but it would seem unlikely that algal cells are present in sufhcient numbers to confer any significant nutritional benefit to the fungus. The morphologically similar Marasmiellus affixus (Berk.) Singer, described from Australia and commonly known as the ‘little stinker’, is also associated with a waxy algae-infected layer. The association of M. affixus with alga was noted by Singer (1973) and has been specu- lated to be a basidio-lichen, although this has not proven (Lepp 2011). A partial, poor quality ITS1 sequence for M. affixus obtained during this work (not deposited) suggests it has affinity with Marasmiellus ramealis (Bull.) Singer rather than the taxa treated here. Acknowledgements Thanks to Dukchul Park, Landcare Research, for DNA extraction and sequencing, and to Sapphire McMullen-Fisher for material of Marasmiellus affixus. The New Zealand De- partment of Conservation is thanked for permission to collect specimens from reserves and national parks that they manage. ‘The first author was supported through the Land- care Research Systematics Portfolio, with Core funding support from the Science and In- novation Group of the New Zealand Ministry of Business, Innovation and Employment. References Antonin V, Noordeloos ME (2010) A Monograph of Marasmioid and Collybioid fungi in Europe, IHW-Verlag. Cooper JA, Leonard PL (2012) Boletopsis nothofagi sp. nov. associated with Nothofagus in the Southern Hemisphere. MycoKeys, 3: 13-22. doi: 10.3897/mycokeys.3.2762 Drummond AJ, Ashton B, Buxton S, Cheung M, Cooper A, Duran C, Field M, Heled J, Kearse M, Markowitz S, Moir R, Stones-Havas S, Sturrock S, Thierer T, Wilson A (2011) Geneious v5.4. Available from: http://www.geneious.com/ 44 Jerry A. Cooper & Pat L. 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