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Mycologia 94(2), 2002, pp. 327-354
© 2002 by The Mycological Society of America

Type studies of sequestrate Russulales II. Australian and New Zealand species related to Russula


Teresa Lebel 1

     Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 97331

Michael A. Castellano

     U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331

   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 LITERATURE CITED
 

Nomenclatural types of the basionyms of species of sequestrate relatives of Russula from Australia and New Zealand were studied and original descriptions emended. Illustrations of microscopic characters are provided for the first time for many species. Several recombinations are proposed, including: (i) Arcangeliella crichtonii comb. nov. ({equiv} Cystangium crichtonii), (ii) Arcangeliella hepaticus comb. nov. ({equiv} Elasmomyces hepaticus), (iii) Macowanites tomentosa comb. nov. ({equiv} Hydnangium tomentosum).

Key words: Basidiomycetes, mycorrhizal, taxonomy, truffle-like Russulales


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 TAXONOMY
 LITERATURE CITED
 
Singer and Smith (1960)Citation distinguished five sequestrate genera related to Russula. Two of these genera, Cystangium and Gymnomyces, were described from Australian specimens. However, prior to 1997, only 12 species of sequestrate relatives of Russula had been described from Australia and New Zealand. The majority of these species are based on collections by Rodway from Tasmania in the 1890s to early 1900s, Cleland from South Australia in the 1910s to 1930s, or Beaton from Victoria in the 1960s to late 1980s (Grgurinovic 1997Citation , Lebel and Castellano 1999Citation ).

Examination of recent collections from Australia and New Zealand revealed a much higher diversity of russuloid sequestrate fungi than previously recognized (Lebel and Castellano 1996Citation , Lebel 1998Citation ). Reexamination of type material was critical to determine relationships among described and new species. The results of our examination of the holotypes of the type species for all seven sequestrate genera of Russulales are presented in Lebel and Trappe (2000)Citation . Following an examination of holotype or isotype material of all sequestrate relatives of Russula described from Australia and New Zealand, complete type descriptions, including many microscopic features not previously described or illustrated, are provided.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
TAXONOMY
 LITERATURE CITED
 
Data on macroscopic features are based on original descriptions and notes accompanying type collections. Colors are in general terms unless stated in original description. Descriptions are emended from an examination of dried specimens to provide complete descriptions in a standardized format. Habitat and fruiting patterns are based on information in original descriptions. Names of herbaria are abbreviated according to Holmgren et al (1990)Citation . A detailed discussion of macroscopic and microscopic characters is presented in Lebel (1998)Citation and Lebel and Trappe (2000)Citation .

Microscopic features observed with a light microscope are described from freehand sections of rehydrated specimens mounted in Melzer's reagent or 5% aqueous KOH (Castellano et al 1989Citation ). Measurements were made at x400 or x1000 with a calibrated optical micrometer. Spore dimensions are given as: length range x width range (mean length ± SE x mean width ± SE, n = 35), and Q ratio. Measurements include neither ornamentation nor the apiculus. Spore shape was determined on the basis of the length-width ratio (Q) for 35 randomly selected spores in 5% aqueous KOH, according to Buyck's (1989)Citation spore shape categories. Scanning electron microscope (SEM) photographs are referred to when available to aid in the interpretation of spore ornamentation patterns (though illustrations and descriptions are in terms of structures visible by light microscopy).

Taxa are presented alphabetically by basionym, with any nomenclatural changes in bold. Illustrations of important characters are given for each species. References to SEM photographs and other studies of type material are listed with the nomenclator for each taxon.


   TAXONOMY
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
LITERATURE CITED
 

Cystangium balpineum Grgur., Larger Fungi of South Australia, 1997. Figs. 17c, d . Figs. 1 , 2 .



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  		 FIG. 17. Cystangium seminudum (= Gymnomyces solidus) A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 1. Cystangium balpineum A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 2. Scanning electron micrographs of spores of Cystangium balpineum. Bars = 10 µm

 
Basidiomata up to 32 mm diam, irregularly globose or flattened with a depressed apex and often indented at base around stipe, exposing gleba for a few mm. Peridial surface dry, smooth, dull ivory white, sometimes with a smoked rosey or purplish rosey tinge. Gleba loculate, the chambers minute, very irregular, elongate, cream-colored or concolorous with the peridium. Stipe up to 6 mm long x 3 mm wide, central, solid, cream-colored, generally not protruding past peridial margin. Columella 1–3 mm wide, percurrent, continuous with stipe, cream colored. Odor absent and taste unknown. Latex absent. Peridiopellis two-layered, a thin suprapellis composed of sparse, repent, branching, hyaline agglutinated hyphae 2–4.5 µm in diam overlying an epithelial subpellis 29–82 µm wide, of 2–4 tiers of irregular, globose, pyriform or elongated inflated cells 16–40 x 12–39 µm; on certain areas of the peridium, the peridiopellis compacted, appearing as a single, agglutinated layer of somewhat inflated elements. Peridial context 110–200 µm wide, heteromerous, mostly of interwoven, hyaline hyphae 2–3.5 µm in diam, with sphaerocysts 14–33 µm in diam in scattered nests. Endocystidia absent. Stipitipellis a cutis 5–14 µm wide, of interwoven, hyaline hyphae 2–4 µm in diam. Stipe-columella context of interwoven, hyaline hyphae 2–4 µm in diam and sphaerocysts 12–32 µm diam in scattered nests. Hymenophoral trama 15–37 µm wide, of interwoven hyphae 2–5.5 µm diam and sphaerocysts 12–32 µm diam, in nests at the intersection of tramal plates; subhymenium 14–23 µm wide, of 1–3 tiers of isodiametric cells 6–14 µm in diam. Clamp connections absent from all tissues. Basidia 22–33.5 x 9–15 µm, hyaline, clavate to broadly clavate, mostly 4-, rarely 2- or 3-sterigmate, sterigmata 5–7 µm long. Cystidia absent. Spores 8.6–11.5 x 7–9.4 µm (9.9 ± 0.32 x 8.1 ± 0.23, n = 35), Q = 1.1–1.2, subglobose to broadly ellipsoid, orthotropic. Ornamentation amyloid, of fairly robust, isolated spines, 0.5–1 µm high. Hilar appendix 1–2.5 x 1–2 µm, central, straight; plage absent. Color of spores in mass white.

Etymology. Derived from the Aboriginal word ‘balpi’, meaning ‘white’, referring to the white color of the peridium.

Specimen examined. AUSTRALIA. SOUTH AUSTRALIA: Mt. Lofty, solitary under litter in Eucalypt forest, 29 Mar 1924, J.B. Cleland, (HOLOTYPE: AD 9757); Mt. Lofty, on ground, 25 May 1924, J.B. Cleland (AD 9755).

Commentary. Cystangium balpineum differs from other described sequestrate Russulales by the combination of the occasional smoked rosey or purplish rosey tints on the peridium, large (16–40 µm) epithelial cells in the peridiopellis, spore ornamentation of low isolated spines and lack of hymenial cystidia. Cystangium sessile also may occasionally have pink to reddish patches on the peridium, however the gleba is sublamellate and the hymenium has abundant, large cystidia rather than a loculate gleba and lacking hymenial cystidia as in C. balpineum.

Cystangium phymatodisporum also resembles C. balpineum. However the basidiomata of C. phymatodisporum lack any pink or red tints, the peridiopellis cells are smaller, 12–22 µm versus 16–40 µm in C. balpineum, the basidia are generally more broadly clavate, spores more broadly ellipsoid with finer, lower ornamentation, and the structure of the hymenium differs from that of C. balpineum.

Cystangium crichtonii G.W. Beaton, Pegler and T.W.K. Young, Trans. Br. Mycol. Soc. 86:181–184. Figs. 1A–D, 2A–E , 1986.

{equiv} Arcangeliella crichtonii (G.W. Beaton, Pegler and T.W.K. Young) T. Lebel et Castellano comb. nov. Figs. 3 , 4 .



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  		 FIG. 3. Arcangeliella crichtonii A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 4. Scanning electron micrographs of spores of Arcangeliella crichtonii. Bars = 10 µm.

 
Basidiomata 15–30 mm in diam, irregularly convex to plane, with a free, undulating margin exposing the lower gleba. Peridial surface pale brown to yellowish brown, discoloring to brown on drying; texture not recorded fresh, dried specimens appearing smooth. Context thin, fragile, creamy white. Gleba pale brown, paler than peridium, sublamellate but strongly anastamosing to appear labyrinthoid-loculate, locules elongate, empty. Stipe 4–6 x 1–2 mm, central to lateral, more or less cylindrical, glabrous, surface concolorous with peridium or slightly paler. Columella percurrent, simple, concolorous with peridium. Odor and taste not recorded. Latex not observed. Peridiopellis two-layered, a thin suprapellis composed of repent, hyaline hyphae 2–6 µm in diam overlying an epithelial subpellis 40–80 µm wide, of 2–4 tiers of irregular, agglutinated, inflated cells 12–22 µm in diam; on certain areas of the peridium, the peridiopellis compacted, appearing as a single, agglutinated layer of somewhat inflated elements. Peridial context 110–180 µm wide, heteromerous, mostly of interwoven, slightly gelatinized, hyaline hyphae 2–3.5 µm in diam, sphaerocysts 10–40 µm in diam in nests, and scattered, sinuous laticiferous hyphae 4–8.5 µm in diam. Endocystidia absent. Stipitipellis a cutis 5–14 µm wide, of interwoven, hyaline hyphae 2–4 µm in diam. Stipe-columella context heteromerous, mostly of interwoven, hyaline hyphae 2–3.5 µm in diam with nested columns of sphaerocysts 12–30 µm in diam. Hymenophoral trama 80–100 µm wide, heteromerous, of interwoven to parallel, hyaline hyphae 2–6 µm in diam, with scattered inflated elements 5–12 µm in diam, sphaerocysts 12–24 µm in diam in nests and laticiferous hyphae 4–8 µm in diam; subhymenium well-developed, 30–45 µm wide, of 2–3 tiers of isodiametric cells 6–14 µm in diam. Clamp connections absent from all tissues. Basidia 36–55 x 8–11 µm, hyaline, cylindrical to ventricose, with 4 sterigmata 4–7 µm long, slightly curved. Cystidia 45–65 x 6–10 µm, hyaline, cylindrical to narrowly ventricose with a tapering, obtuse apex and some granular contents; generally not projecting beyond basidia, arising in the trama, rare. Spores 8–10 x 7–9.5 µm (9.2 ± 0.4 x 8.5 ± 0.3, n = 35), Q = 1.09–1.18, subglobose to broadly ovoid, heterotropic, asymmetric, hyaline. Ornamentation strongly amyloid, of interrupted and anastamosing concentric ridges, at times radial or random, up to 1 µm high. Hilar appendix hyaline, eccentric, conical, 1–1.5 x 1 µm; plage absent or when present, inconspicuous, inamyloid. Color of spores in mass not recorded for fresh material, off-white to cream-colored when dried.

Etymology. After the Australian collector George Crichton.

Specimen examined. AUSTRALIA. NEW SOUTH WALES: Tuncurry, epigeous, scattered or caespitose, under Eucalyptus debris, 9 August 1984, G. Crichton, Crichton Z47 (HOLOTYPE: K).

Commentary. An emphasis on presence of laticiferous hyphae versus latex production and of pseudocystidia as delimiting characters between Russula and Lactarius lineages will affect where this species is placed. Beaton et al (1986)Citation point out the similarities between Zelleromyces striatus and this species, but argue ingenuously that as Arcangeliella has not been found in Australia, this species should be placed in Cystangium rather than the equivalent genus of the Lactarius lineage. However, recent work suggests that Arcangeliella is more common in Australia than previously recognized (J. Trappe pers comm).

The general morphology and texture of the basidiomes, presence of laticiferous hyphae in the peridial context and hymenophoral trama, presence of pseudocystidia, and spore ornamentation type (concentric ridges) all point to a closer relationship of this taxon with Lactarius rather than with Russula. The epithelial peridiopellis in this case is of less importance than the presence of abundant laticiferous hyphae in determining the placement of this species in Arcangeliella rather than Cystangium. Care should be taken as specimens of Arcangeliella collected in dry weather or not cut and examined in the field at time of collection often do not exude latex (Thiers 1984a, bCitation ). The only other Arcangeliella species described from Australia, A. texta, differs in several features, particularly its mostly isolated spore ornamentation rather than concentric ridges and trichodermial rather than epithelial peridiopellis (Smith 1962Citation ).

Cystangium phymatodisporum G.W. Beaton, Pegler and T.W.K. Young, Kew Bull. 39:672–674. Fig. 3E–G , pl 24K–P, 1984. Figs. 5 , 6 .



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  		 FIG. 5. Cystangium phymatodisporum A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 6. Scanning electron micrographs of spores of Cystangium phymatodisporum. Bars = 10 µm.

 
Basidiomata 10–30 mm in diam, broadly ellipsoid, depressed at apex and around stipe, margin strongly incurved and completely covering gleba. Peridial surface smooth and glabrous or finely pruinose, dry, becoming wrinkled on drying, white, staining brown. Context thin, fragile, off-white when dry. Gleba white becoming ivory, sublamellate to loculate, locules labyrinthine, elongated, irregular, 0.5–1 mm in diam. Stipe 2–5 x 1–2 mm, short, curved, not projecting beyond peridium, solid, surface white. Columella percurrent, simple, narrow, white, sometimes free from the gleba. Odor and taste not recorded. Latex absent. Peridiopellis two-layered, a thin, inconspicuous suprapellis of repent, interwoven, agglutinated hyaline hyphae 2–3 µm in diam, with some inflated hyphal tips, clavate to globose 7–13 x 4–8 µm, overlying a well-developed epithelial subpellis 35–90 µm broad composed of inflated cells 12–22 µm in diam. Peridial context 50–80 µm wide, of interwoven to subparallel, hyaline hyphae 2–3 µm in diam. Endocystidia and oleiferous hyphae absent. Stipitipellis undifferentiated from context. Stipe-columella context heteromerous, of interwoven, hyaline hyphae 1.5–3 µm in diam and sphaerocysts 14–32 µm in diam in nests. Hymenophoral trama 30–65 µm wide, heteromerous, with narrow, ± parallel, hyaline hyphae 2–3.5 µm in diam, and sphaerocysts 14–26 µm in diam in scattered nests; subhymenium well-developed, 18–25 µm wide, with 2–3 tiers of ± isodiametric cells 5–14 x 4–12 µm. Clamp connections absent from all tissues. Basidia 15–38 x 10–14 µm, hyaline, broadly clavate to clavate, with 4 sterigmata 3–6 x 1–1.5 µm. Cystidia absent. Spores 8–10 x 7–8.5 µm (8.9 ± 0.3 x 7.1 ± 0.2, n = 35), Q = 1.17–1.25, broadly ellipsoid, suborthotropic and symmetric, hyaline. Ornamentation amyloid, of crowded somewhat irregularly shaped, isolated warts 0.5–0.8 µm high. Hilar appendix prominent, 1–2 x 0.8–1.5 µm, central, truncate; plage inconspicuous, inamyloid. Spores white in mass.

Etymology. Derived from the Latin terms phymatodeus and sporum, referring to the warty or verrucose spore ornamentation.

Specimen examined. AUSTRALIA. VICTORIA: Eildon State Park, Dry Creek, emergent under Eucalyptus sp., 14 August 1982, C. Beauglehole, K. and G. Beaton, Beaton 50 (HOLOTYPE: K).

Commentary. Beaton et al (1984)Citation differentiated this species from Cystangium sessile on the basis of smaller, more ellipsoid spores bearing a lower ornamentation of isolated warts. Other microscopic differences include the absence of hymenial cystidia and lack of sphaerocysts in the peridial context of C. phymatodisporum. Macroscopically, C. phymatodisporum has a loculate gleba rather than a sublamellate gleba as in C. sessile. Cystangium balpineum resembles C. phymatodisporum in the lack of hymenial cystidia and broadly ellipsoid spores with a low ornamentation of isolated warts. However, C. balpineum basidiomata may have smoked rosey to purplish patches on the peridium which C. phymatodisporum lacks, the peridiopellis cells are generally larger though in fewer tiers, and the structure of the hymenium differs from that of C. phymatodisporum.

Elasmomyces hepaticus G.W. Beaton, Pegler and T.W.K. Young, Kew Bull. 39 (4):676. Fig. 4A–C , pl 25E–J, 1984.

{equiv} Arcangeliella hepaticus (G.W. Beaton, Pegler and T.W.K. Young) T. Lebel et Castellano comb. nov. Figs. 7 , 8 .



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  		 FIG. 7. Arcangeliella hepaticus A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 8. Scanning electron micrographs of spores of Arcangeliella hepaticus. Bars = 10 µm.

 
Basidiomata 4–26 mm in diam, subglobose to irregularly ellipsoid, depressed at the base with an exposed gleba and short stipe. Peridial surface dry, smooth becoming wrinkled, dark reddish brown. Gleba pale orange, loculate to sublamellate, locules elongated, 1–2 mm in diam. Stipe 1–3 x 0.5–1 mm, curved, cylindrical, solid, surface and context dark brown, not protruding much past peridium at base. Columella percurrent, simple or weakly dendroid, narrow, dark brown. Odor and taste not recorded. Latex absent. Peridiopellis 40–80 µm wide, an epithelium, composed of 3–6 tiers of agglutinated, brownish inflated cells 12–39 µm in diam. Peridial context 80–110 µm wide of densely packed, subgelatinous, interwoven, hyaline hyphae 2–3 µm in diam, with scattered, sinuous, laticiferous hyphae 3–7 µm in diam; sphaerocysts absent from context. Endocystidia and pileocystidia absent. Stipitipellis undifferentiated from context. Stipe-columella context heteromerous, of interwoven, hyaline hyphae 2–3 µm in diam, sphaerocysts 16–27 µm in diam in nests, and scattered, sinuous, laticiferous hyphae 4–7 µm in diam. Hymenophoral trama 30–65 µm wide, of densely packed, interwoven, subgelatinous, hyaline hyphae 2–5.5 µm in diam, with a few scattered, sinuous, laticiferous hyphae 3–7 µm in diam extending into the subhymenium; sphaerocysts absent; subhymenium well-developed, 25–38 µm broad, with 2–3 tiers of isodiametric cells 5–12 µm in diam. Clamp connections absent from all tissues. Basidia 36–52 x 7–10.5 µm, hyaline, cylindrical to clavate, with 4 slender sterigmata 4–10 µm long. Cystidia absent. Spores 6.5–9 x 6–8 µm (8.1 ± 0.3 x 7.4 ± 0.3, n = 35), Q = 1.02–1.07, globose to subglobose, orthotropic or asymmetric, hyaline. Ornamentation strongly amyloid, of irregular ridges 0.8–1.5 µm high, forming a fine complete reticulum. Hilar appendix 1–2 x 0.5–1 µm, cylindrical, eccentric; plage inconspicuous, inamyloid. Spores white in mass.

Etymology. Derived from the latin word ‘hepaticus’, meaning ‘liver-colored’, referring to the color of the peridium.

Specimen examined. AUSTRALIA. VICTORIA: Black Range Rd., near Buxton, hypogeous under mixed Eucalyptus spp., 27 May 1980, K and G. Beaton, Beaton 69 (HOLOTYPE: K).

Commentary. Beaton et al (1984)Citation emphasized the exposed basal gleba, symmetric spores, lack of sphaerocysts in the hymenophoral trama, and absence of latex production, to place this species in Elasmomyces. An emphasis on presence of laticiferous hyphae versus latex production as a delimiting character between Russula and Lactarius affects the placement of Arcangeliella hepaticus. The reticulate spore ornamentation, lack of sphaerocysts in the hymenophoral trama and peridial context, presence of laticiferous hyphae, structure of the peridiopellis, and general form and color of basidiomata all point to the close relationship of this species with Lactarius rather than with Russula.

Arcangeliella hepaticus differs from A. crichtonii in that the latter has radially oriented ridges on the spores and a pale brown peridium versus completely reticulate spore ornamentation and a dark brown peridium. Arcangeliella texta has a mostly isolated rather than a completely reticulate spore ornamentation as in A. hepaticus (Smith 1962Citation ).

Gymnomyces eildonensis G.W. Beaton, Pegler and T.W.K. Young, Kew Bull. 39 (4):680. Fig. 5E–H , pl 26F–L, 1984. Figs. 9 , 10 .



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  		 FIG. 9. Gymnomyces eildonensis A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 10. Scanning electron micrographs of spores of Gymnomyces eildonensis. Bars = 10 µm.

 
Basidiomata 15–30 mm in diam, subglobose or ellipsoid, sometimes contorted and becoming lobed, basally depressed around the point of attachment. Peridial surface glabrous, becoming wrinkled, sometimes cracking and breaking away, at first cream colored to pale ochraceous, drying dark red-brown, with red-yellow patches readily visible in protected areas. Gleba cream colored drying grayish orange, loculate, locules small, round or elongated, 1.5–4 mm long, sometimes with random concentric or radial arrangement. Stipe absent or present as a rudimentary sterile base, white, small. Columella absent. Odor and taste not recorded. Latex absent. Peridiopellis either appearing as an ixocutis 8–14 µm wide, or in patches as an agglutinated turf 10–26 µm wide, of mostly repent, hyaline to pale brown hyphal tips 2–3 µm in diam. Peridial context 80–350 µm wide, of interwoven, hyaline hyphae 2–3.5 µm in diam, with abundant straight to sinuous, refractive hyphae 2–7 µm in diam, some extending into the peridiopellis, and nests of sphaerocysts 14–32 µm in diam, becoming more common near the subhymenium. Endocystidia absent. Hymenophoral trama 12–32 µm wide, heteromerous, of interwoven, hyaline hyphae 2–4 µm in diam, scattered inflated elements 4–6 µm in diam, refractive hyphae 2–6 µm in diam, and abundant sphaerocysts 12–28 µm in diam in nests; subhymenium well-developed, 15–30 µm wide, with 1–3 tiers of ± isodiametric cells 6–10 µm in diam and scattered inflated cells 12–22 µm in diam. Clamp connections absent from all tissues. Basidia 28–42 x 8–12 µm, hyaline, clavate, with 2 and 4 sterigmata 3–6 x 1–1.5 µm. Cystidia 32–54 x 6–11 µm, fusoid-ventricose, apex mucronate or obtuse, with some refractive granular contents in KOH; arising in trama, not extending beyond basidia, scattered. Spores 7–9 x 7–8.5 µm (8.1 ± 0.34 x 7.6 ± 0.4, n = 35), Q = 1.06–1.11, subglobose, orthotropic and symmetrical, hyaline. Ornamentation incompletely amyloid, of abundant, irregular warts and truncate rods 0.5–0.8 µm high, isolated or connected at the base to neighbors to form short, branched, irregular ridges <0.3 µm high in a poorly-developed partial reticulum. Examination of SEM photos shows the warts and ridges to be connected in an almost complete reticulum, which is not readily apparent with the light microscope. The amyloid covering of the ornamentation is apparently incomplete, covering mostly the apices of warts and ridges. Hilar appendix prominent, 1–2 x 1–1.5 µm, tapering; plage absent. Spores white to pale tan in mass.

Etymology. Derived from the location of the type collection, ‘Eildon.’

Specimen examined. AUSTRALIA. VICTORIA: Junction of Snobs Creek and Conns Gap near Eildon, under mixed Eucalyptus, 3 May 1982, K. and G. Beaton, Beaton 2 (ISOTYPE: K).

Commentary. Gymnomyces eildonensis resembles G. pallidus. The peridiopellis of Gymnomyces pallidus is a turf of interwoven hyphae and cystidia, which may be agglutinated in some specimens, appearing similar to the ixocutis and patchy turf of brown-walled hyphae of G. eildonensis. Spore ornamentation in G. eildonensis is quite different, though the differences may be difficult to discern by light microscopy. Gymnomyces eildonensis spore ornamentation is generally lower, 0.3–0.8 µm versus 0.5–1 µm, appears denser with more connections between elements, and the spores are slightly smaller than those of G. pallidus. Beaton et al (1984)Citation do not mention the gleba darkening to sordid white in G. eildonensis as it does in G. pallidus (Massee and Rodway 1898Citation ), but they do mention the darkening of the peridium in both species. Gymnomyces eildonensis has distinct red tones to the peridium and red-yellow patches readily visible in protected areas unlike G. pallidus in which the peridium becomes overall sordid brown with no apparent red tones.

Gymnomyces megasporus Rodway, Paps. & Proc. Roy. Soc. Tasmania 1925:168. 1926.

{equiv} Octaviania megaspora (Rodway) G. Cunn., New Zealand J. Sci. Technol. 22:300B. 1941. pl 33, fig. 18.

{equiv} Cystangium megasporum (Rodway) T. Lebel et Castellano comb. nov. Figs. 11 , 12 .



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  		 FIG. 11. Cystangium megasporum A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 12. Scanning electron micrographs of spores of Cystangium megasporum. Bars = 10 µm.

 
Basidiomata 10–15 mm in diam, depressed globose. Peridial surface smooth to irregularly roughened, pallid cream to brown. Gleba cream colored, loculate, locules small, irregular, dense. Stipe absent. Columella absent. Odor and taste not recorded. Latex absent. Peridiopellis two-layered, composed of a suprapellis 10–28 µm wide, of tangled, mostly repent, hyaline hyphae 2–3 µm in diam overlying an epithelial subpellis 45–80 µm wide, of 4–6 tiers of inflated cells 8–19 x 6–19 µm in diam. Peridial context 30–50 µm wide, of interwoven, hyaline hyphae 2–3.5 µm in diam, with sphaerocysts 14–28 µm in diam in scattered nests. Endocystidia absent. Hymenophoral trama 30–50 µm wide, of compact, interwoven, subgelatinous, hyaline hyphae 2–3.5 µm in diam and sphaerocysts 12–28 µm in diam in scattered nests; subhymenium not rehydrating well, appearing to be 16–26 µm wide, with 2–3 tiers of ± isodiametric cells 4–11 x 5–10 µm. Clamp connections absent from all tissues. Basidia 28–42 x 9–12 µm, hyaline, cylindrical to narrow clavate, mostly with 1 or rarely 2 robust sterigmata, 6–11 x 1–2.5 µm. Cystidia absent. Spores 12–16.5 x 11.5–16 µm (14.8 ± 0.2 x 14.4 ± 0.4, n = 35), Q = 1.01–1.03, globose, orthotropic, hyaline. Ornamentation strongly amyloid, on most spores of dense, robust, irregular, isolated warts, rods and wedges, 2–3 µm high x 1–2 µm wide; on a few spores the ornamentation is of very dense, isolated warts and rods, fairly similar in appearance, ±1 µm high. Elements appearing irregular, hemispherical, rod-like in surface view. Hilar appendix inconspicuous, 0.5–1 x 1 µm; plage absent. Spore color in mass unknown fresh, appearing white when dried.

Etymology. Derived from the Greek word ‘megas’, meaning ‘large’, referring to the large spores.

Specimen examined. AUSTRALIA. TASMANIA: Cascades, under Eucalyptus sp., May 1925, L. Rodway, HO 113639 (HOLOTYPE: HO). TASMANIA: 8638 (ISOTYPE: PDD).

Commentary. Rodway (1926)Citation distinguished Cystangium megasporum from Gymnomyces pallidus and C. seminudum by the larger spores. Cunningham (1941)Citation apparently being unaware of the iodine reaction of spore ornamentation in the Russulales, placed C. megasporum in Octaviania because of the robust, isolated spore ornamentation. Following the revision of generic boundaries in the sequestrate Russulales, Gymnomyces is restricted to species lacking an epithelial peridiopellis (Lebel and Trappe 2000Citation ). The presence of an epithelial peridiopellis, isolated spore ornamentation, and general appearance of the fruitbody, place this species in the genus Cystangium (Lebel and Trappe 2000Citation ).

Cystangium megasporum resembles C. seminudum macroscopically and microscopically, differing mainly in the larger spores, 12–16 µm versus 9–12 µm and the very robust, irregular appearance and dense nature of the spore ornamentation. The basidia of Cystangium megasporum generally have 1 or 2 sterigmata and hymenophoral cystidia were not observed, whereas C. seminudum has basidia with 2 or 4 sterigmata and prominent hymenophoral cystidia. Variation in sterigmata number is known to affect spore size, basidia with 1 and 2 sterigmata often have larger spores than basidia with 4 sterigmata. However in this particular case, larger spores of Cystangium seminudum never have the robust wedges and warts of C. megasporum spores.

Gymnomyces pallidus Massee and Rodway, in Massee, Kew Bull. Misc. Inform. 1898:125. 1898.

{equiv} Octaviania pallida (Massee and Rodway) G. Cunn., Proc. Linn. Soc. New South Wales 60:119. 1935. pl 33, Fig. 17 .

Illustrations of type material. Pegler and Young (1979)Citation ; Lebel and Trappe (2000)Citation . Figs. 13 , 14 .



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  		 FIG. 13. Gymnomyces pallidus A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Pileicystidia. G. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 14. Scanning electron micrographs of spores of Gymnomyces pallidus. Bars = 10 µm.

 
Specimens examined. AUSTRALIA. TASMANIA: West coast, Jun 1896, L. Rodway, Rodway 299 (HOLOTYPE: K); 113643 (ISOTYPE: HO); Collected June 1920, Rodway 111 (ISOTYPE: BPI).

Etymology. Derived from the latin term ‘pallidus’, meaning ‘pale’, referring to the white to cream colored gleba.

Commentary. A discussion of the taxonomic and nomenclatural history, and a full description of the holotype of Gymnomyces pallidus can be found in Lebel and Trappe (2000)Citation . Gymnomyces pallidus can be distinguished from other species of Gymnomyces by the irregular, incompletely amlyoid, 0.5–1 µm high spore ornamentation, scattered refractive hymenial cystidia, and change in color of the peridium from white to sordid brown.

Dr. T. May (pers comm) considered the writing on the two isotype packets examined to be that of Rodway. Rodway's use of ‘type’ and ‘isotype’ was rather inconsistent, and both of these collections were made after the type was described. ISOTYPE. 113643 (HO) includes 2 collections. (1) is Cystangium seminudum: white, globose, spores with dense amyloid ornamentation of isolated spines 2–3 µm high, cystidia abundant, large; peridiopellis two-layered, subpellis an epithelium with overlying repent hyphal suprapellis; (2) basidioma brown-yellow; spores with amyloid, reticulate ornamentation ±0.8 µm high. Does not match any described species of Gymnomyces. ISOTYPE. Rodway 111 (BPI). Collected June 1920. 2 collections, (1) is G. pallidus: brownish, spore ornamentation of irregular isolated elements, irregularly amyloid; peridiopellis a dense turf of upright cystidia and hyphae; (2) is Cystangium seminudum: white, globose; spores with dense, amyloid, isolated spines ±3 µm high; peridiopellis 2-layered.

Gymnomyces seminudus Massee and Rodway in Massee, Kew Bull. Misc. Inform. 1898:125. 1898.

{equiv} Arcangeliella seminuda (Massee and Rodway) Zeller and CW. Dodge, Ann. Missouri Bot. Gard. 23:617. 1937.

{equiv} Octaviania seminuda (Massee and Rodway) G. Cunn., Trans. Proc. Roy. Soc. New Zealand 67:408. pl 33, Fig. 19 , 1938.



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  		 FIG. 19. Gymnomyces wirrabarensis A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 
{equiv} Cystangium seminudum (Massee and Rodway) T. Lebel et Castellano comb. nov.

Illustrations of type material. Pegler and Young (1979)Citation , Beaton et al (1984)Citation . Figs. 15 , 16 .



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  		 FIG. 15. Cystangium seminudum A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 16. Scanning electron micrographs of spores of Cystangium seminudum. Bars = 10 µm.

 
Basidiomata 15–35 mm in diam, globose to subglobose, gleba completely enclosed. Peridial surface smooth to delicately tomentose, dry, white becoming ochre. Context fragile, white. Gleba white, loculate, locules compressed, small, irregular. Stipe absent. Columella absent or rudimentary, narrow, simple. Odor and taste not recorded. Latex not recorded. Peridiopellis two-layered, composed of a narrow suprapellis 20–70 µm wide, of tangled hyphae, repent or erect, 25–70 x 3–5 µm, overlying an epithelial subpellis 28–55 µm wide, composed of 3–5 inflated cells 8–17 x 8–20 µm in diam. Peridial context 28–44 µm wide, of interwoven, hyaline hyphae 2–3.5 µm in diam with sphaerocysts 12–26 µm in diam in scattered to abundant nests. Endocystidia absent. Stipitipellis absent. Columella context when present, of interwoven, hyaline hyphae 2–3 µm in diam. Hymenophoral trama 35–65 µm wide, heteromerous, composed of hyaline hyphae 2–4 µm in diam and abundant sphaerocysts 15–25 µm in diam in nests; subhymenium well-developed, 12–20 µm wide, with 2–4 tiers of isodiametric cells 4–12 x 4–10 µm. Clamp connections absent from all tissues. Basidia 28–52 x 7–11 µm, hyaline, ventricose to clavate, mostly with 2 or rarely 4 robust sterigmata 5–10 x 1–2.5 µm. Cystidia 52–100 x 8–15 µm, hyaline with few granular contents refractive in KOH, cylindrical to clavate with obtuse apices; extending beyond basidia considerably, arising at the base of the subhymenium, scattered to abundant. Spores 10–12.5 x 9.5–12 µm (11.2 ± 0.7 x 10.5 ± 0.6, n = 35), Q = 1.05–1.08, globose, orthotropic, symmetric, hyaline. Ornamentation amyloid, of robust, crowded, laterally compressed spines 1.5–2.5 x 1–1.5 µm, some coalescing at bases. Hilar appendix 2–2.5 x 0.5–1 µm, cylindrical; plage inamyloid. Spores in mass white.

Etymology. Derived from the latin term seminudus, meaning ‘partially naked’, referring to the apparently rudimentary peridium.

Specimen examined: AUSTRALIA. TASMANIA: hypogeous to emergent, L. Rodway, Rodway 124 (HOLOTYPE: HO; ISOTYPE: FH).

Commentary. Massee and Rodway (1898)Citation described two species of Gymnomyces, G. pallidus and G. seminudus. They distinguished Gymnomyces from Octaviania based on the apparent absence of a peridium on the former. Cunningham (1938)Citation transferred both species to Octaviania, apparently unaware of the iodine reaction of the spore ornamentation of the Russulales. Zeller and Dodge (1937)Citation transferred G. seminudus to Arcangeliella, as they felt this genus was "carefully and fully described," while Gymnomyces was not well supported. Singer and Smith (1960)Citation subsequently maintained the name Gymnomyces as a well-differentiated group. However, the presence of an epithelial peridiopellis, large hymenial cystidia, and isolated spore ornamentation, places this species alongside Cystangium megasporum, in an expanded Cystangium (Lebel and Trappe 2000Citation ).

Cystangium seminudum differs from other species of Cystangium microscopically in the combination of a suprapellis of tangled hyphae, prominent hymenial cystidia, and spores ornamented with isolated spines 1.5–2.5 x 1–1.5 µm; macroscopically the smooth, generally white to creamy peridium and lack of a stipe is distinctive. The Rodway 124 (H0113648) packet had four different envelopes enclosed: (1) is 124 HOLOTYPE; (2) is 124 cotype. "G. seminudus" Collected August 1898, Australia, Tasmania. White, globose, smooth; spore ornamentation isolated spinose, peridiopellis two-layered. Matches the holotype of Cystangium seminudum; (3) 124 "G. seminudus." Spores larger than the holotype, ±14 µm, ornamentation less dense, spines more irregular, curved, acute. Peridiopellis two-layered, hymenial cystidia present, large. An unknown Cystangium sp.; (4) 124 COTYPE postal envelope. Does not match C. seminudum, or #3. Another unknown Cystangium sp.

Gymnomyces solidus Rodway, Paps. & Proc. Roy. Soc. Tasmania 1920:157. 1921.

= Cystangium seminudum (Massee and Rodway) T. Lebel et Castellano comb. nov. Figs. 17 , 18 .



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  		 FIG. 18. Scanning electron micrographs of spores of Cystangium seminudum (G. solidus). Bars = 10 µm.

 
Basidiomata 10 mm in diam, irregularly globose. Peridial surface none, the tramal plates defining the glebal locules protruding externally. Gleba white, loculate, of closely packed locules 0.3 mm in diam, full of spores. Stipe or columella not recorded, none apparent in specimens examined. Odor and taste not recorded. Latex not recorded. Peridiopellis apparently lacking on portion of holotype examined. Peridial context 35–70 µm wide, of interwoven, hyaline hyphae 2–3 µm in diam; sphaerocysts absent. Endocystidia absent. Hymenophoral trama 22–37 µm wide, of compacted, interwoven, hyaline hyphae 2–3 µm in diam (not rehydrating well) and sphaerocysts 12–24 µm in diam in scattered to abundant nests; subhymenium 15–25 µm wide, with 1–2 tiers of isodiametric cells 6–14 x 6–12 µm. Clamp connections absent from all tissues. Basidia 28–39 x 7–10.5 µm, clavate to cylindrical, hyaline, at least some with 2 sterigmata but mostly with 4, 4–7 µm long. Cystidia 40–62 x 9–12 µm, hyaline, clavate to ventricose with an obtuse apex; extending beyond basidia, arising at the base of the subhymenium, abundant. Spores 8.5–10.5 x 8.5–10 µm (9.6 ± 0.4 µm x 9.3 ± 0.3 µm, n = 35), Q = 1.01–1.04, globose, orthotropic, symmetric, hyaline. Ornamentation amyloid, of robust, somewhat curved, isolated spines and warts, some appearing laterally compressed, 2–3 x 1 µm. Elements distributed fairly coarsely, of regular height and amyloid reaction. Hilar appendix small, central, 1–1.5 x 1 µm; plage absent. Spore white in mass.

Etymology. Derived from the latin word solidus, meaning ‘solid,’ referring to the closely packed locules of the gleba.

Specimen examined: AUSTRALIA. TASMANIA: Cascades, Mt Wellington, July 1920, L. Rodway, HO 113647 (ISOTYPE: HO).

Commentary. Not much information is available on this species as it hasn't been studied since originally described. Rodway's (1921)Citation diagnosis lacks much in the way of macroscopic characters, which causes some difficulty as the peridiopellis appears to have been lost. However, the spore ornamentation, basidia and cystidia shape and size, and hymenial structure are consistent with a young Cystangium seminudum. We propose that G. solidus be synonymized with C. seminudum.

Gymnomyces wirrabarensis Grgur., Larger Fungi of South Australia, 1997. Fig. 37 Figs. 19 , 20 .



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  		 FIG. 20. Scanning electron micrographs of spores of Gymnomyces wirrabarensis. Bars = 10 µm.

 
Basidiomata 12–30 mm diam, subglobose or flattened globose. Peridial surface smooth, dry, sometimes almost wanting, white to pallid white becoming ochraceous. Gleba loculate, the chambers 0.5–1 mm diam, labyrinthiform, white becoming pale ochraceous. Stipe absent. Columella absent or rudimentary if present, ±1 mm wide, concolorous or slightly paler than gleba. Odor and taste not recorded. Latex absent. Peridiopellis. 7–32 µm wide, a cutis of mostly repent, interwoven, hyaline hyphae 2–4 µm diam; the hyphal tips sometimes projecting in younger specimens as a patchily upright turf. Peridial context 75–205 µm thick, of interwoven, somewhat agglutinated hyphae, 2.5–6.5 µm diam. Endocystidia absent. Stipitipellis absent. Hymenophoral trama 24–37 µm wide, heteromerous, of hyaline, interwoven hyphae 2–4.5 µm diam and sphaerocysts 14–26 µm diam in scattered nests in junctions of trama; subhymenium 10–19 µm wide, not well-developed, ramose. Clamp connections absent from all tissues. Basidia 33–49.5 x 8.5–13 µm, cylindrical to narrowly clavate, 2-, 3- or 4-sterigmate; sterigmata 6–12.5 µm long. Cystidia 32–56 x 5–9.5 µm, cylindrical to irregularly fusoid, the apices obtuse or sometimes capitate, with some granular contents; rare, arising in the hymenium. Spores 8–12 x 7.5–11.5 µm (10.2 ± 0.47 x 9.9 ± 0.51, n = 35), Q = 1.05–1.12, globose to subglobose, orthotropic to suborthotropic. Ornamentation amyloid, of numerous warts or short spines often joined by short basal connectives to form a partial or almost complete reticulum, ±0.5 µm high. Hilar appendix 1–2 x 1–2 µm, central; plage absent. Spores white in mass.

Etymology. Derived from the Aboriginal word wirrabara, meaning ‘tree place’.

Specimen examined. AUSTRALIA. SOUTH AUSTRALIA: Mt. Lofty, solitary or in groups in Eucalyptus forest, 18 Jun 1932, J.B. Cleland, (HOLOTYPE: AD 5843).

Other specimen: AD9720 South Australia: Belair National Park 26 June 1932, J.B. Cleland.

Commentary. Gymnomyces wirrabarensis may be distinguished from other species of sequestrate Russulales by the globose to subglobose spores ornamented with a partial to almost complete reticulum, undifferentiated peridiopellis and long, cylindrical to narrowly clavate basidia.

Hydnangium tomentosum J.W. Cribb, Univ. Queensland Dept. Bot. Pap. 3:251. Fig. 9 . 1958.

{equiv} Martellia tomentosa (J.W. Cribb) A.H. Sm., Mycologia 54:631. 1962.

{equiv} Macowanites tomentosa (J.W. Cribb) T. Lebel et Castellano comb. nov.

Illustrations of type material. Pegler and Young (1979)Citation . Figs. 21 , 22 .



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  		 FIG. 21. Macowanites tomentosa A. Basidioma. B. Hymenophoral trama and hymenium. C. Peridiopellis and peridial context. D. Basidia. E. Hymenophoral cystidia. F. Peridiopellis cystidia. G. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 22. Scanning electron micrographs of spores of Macowanites tomentosa. Bars = 10 µm.

 
Basidiomata up to 15 mm in diam, subglobose. Peridial surface tomentose, ochraceous. Gleba white becoming pale brown, loculate, locules labyrinthiform, up to 1 mm in diam, empty. Stipe either 1–5 x 1–2 mm, solid, off-white, vestigial, or reduced to basal pad. Columella absent or present, rudimentary, concolorous with gleba. Odor and taste not recorded. Latex not recorded. Peridiopellis 10–90 µm wide, a tangled trichodermium of upright to collapsed, septate, rarely branched, hyaline to brown hyphae 12–32 x 3–6 µm, the terminal cells sometimes inflated, fusoid or clavate with obtuse or mucronate apices, with pigmented granular contents orange brown in KOH which may be agglutinated, repent in older specimens, and an underlying gelatinized layer, 15–25 µm wide, of hyaline hyphae 2–4 µm in diam, the cells appearing ± isodiametric. In younger basidiomata, the peridiopellis a hymeniform turf of tightly packed clavate to cylindrical cells 25–35 x 3–7 µm, with an overlying gelatinous layer 12–18 µm thick. Peridial context 70–200 µm wide, of interwoven, hyaline to pale brown hyphae 2–3.5 µm in diam, with abundant straight to sinuous, refractive hyphae 2–7 µm in diam, some of which extend into the peridiopellis, and scattered nests of sphaerocysts 12–25 µm in diam. Endocystidia absent. Stipitipellis 9–15 µm wide, a turf of hyphal tips 2–4 µm in diam and scattered, inflated, fusoid or ventricose cystidia 8–11 x 3–6 µm, with obtuse or acute apices. Columella context when present, of interwoven, hyaline hyphae 2–3 µm in diam and sphaerocysts 12–22 µm in diam in scattered nests. Hymenophoral trama 35–82 µm wide, of densely packed, interwoven, hyaline hyphae 2–4 µm in diam, scattered inflated elements 4–6 µm in diam, sinuous refractive hyphae 2–6 µm in diam and a few sphaerocysts 13–27 µm in diam in scattered nests; subhymenium not rehydrating well, 14–26 µm wide, with 1–2 tiers of isodiametric cells 6–10 µm in diam. Clamp connections absent from all tissues. Basidia 24–36 x 7–13 µm, hyaline, cylindrical to clavate, with 4 (-2) sterigmata 5–7 x 1–1.5 µm (not rehydrating well). Cystidia dimorphic, the most abundant 22–34 x 4–9 µm, cylindrical or fusoid-ventricose with mucronate or obtuse apices and some refractive granular content or oily contents golden in KOH, arising in trama and subhymenium, but rarely extending beyond basidia; rarer, 29–80 x 10–18 µm, clavate or cylindrical with obtuse apices and some granular refractive contents, arising in the subhymenium, protruding well past basidia. Spores 8.5–10 x 8–9.5 µm (9.4 ± 0.25 x 8.8 ± 0.5, n = 35), Q = 1.04–1.08, globose to subglobose, orthotropic and symmetrical, hyaline. Ornamentation irregularly amyloid, of abundant, isolated warts and truncate rods or spines, 0.5–1 µm high, irregular in shape, size and height, often slightly curved, rarely the bases of 1–2 elements coalescing. Hilar appendix 1–1.5 x 1–1.5 µm, central, tapering; plage absent. Spores in mass off-white to pale brown when dried.

Etymology. Derived from the latin word tomentosus, meaning ‘densely interwoven hair covering’, referring to the tomentose peridium.

Specimen examined: AUSTRALIA. QUEENSLAND: Lamington National Park, "in leaf mould", 22 March 1952, J. Herbert, PDD 12329 (ISOTYPE: PDD).

Commentary. Cribb (1958)Citation tended to follow Cunningham in placing all the round spored species of the Hymenogastraceae in either Octaviania or Hydnangium. Smith (1962)Citation transferred this species to Martellia because of the amyloid reaction of the spore ornamentation. However, Macowanites tomentosa is considered to be a member of the genus Macowanites and not a Gymnomyces as it consistently has a stipe-columella, more complicated peridiopellis, and hymenophoral cystidia (Lebel and Trappe 2000Citation ).

Microscopically this collection resembles Gymnomyces pallidus. The spore ornamentation of Macowanites tomentosa closely matches that of G. pallidus, though the warts are more abundant with fewer connections between elements, and the spores are, on average, slightly larger. Macowanites tomentosa has a distinctly filamentous peridiopellis and the hymenophoral cystidia are dimorphic, not a short turf of hyphal tips and cystidia and monomorphic hymenophoral cystidia as in Gymnomyces pallidus. Macroscopically these two species differ in the presence of a stipe-columella in M. tomentosa and lack of such in G. pallidus.

Macowanites carmineus R.F. McNabb, New Zealand J. Bot. 9:359. Fig. 1 . m–o, 1971. Figs. 23 , 24 .



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  		 FIG. 23. Macowanites carmineus A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 24. Scanning electron micrographs of spores of Macowanites carmineus. Bars = 10 µm.

 
Basidiomata 30–60 mm in diam, hemispherical to depressed-globose with strongly involute margins when young, centrally depressed with involute margins at maturity, stipitate. Pileal surface slightly viscid, glabrous to slightly pruinose, bright carmine red to dark red, red pigment leaching under wet conditions. Context white, firm, unchanging with age. Gleba chalk white, sublamellate, attached to apex of stipe-columella, convoluted, glebal plates fragile, radiating from stipe-columella to margin, forming narrow locules exposed at lower margin of pileus. Stipe 30–55 x 14–18 mm, more or less equal, or slightly tapered at base, solid to hollowed, dry, faintly longitudinally striate, surface and context chalk white. Columella percurrent, simple, white. Odor and taste mild. Latex absent. Chemical tests: formalin on context no reaction; phenol on context slowly deep vinaceous; FeSO4 on context no reaction; guaicol on stipe-columella base no reaction; KOH on peridium bleaching action on red color leaving pallid orange area, on context faint yellow; NH4OH on peridium faint purplish flush, on context no reaction. Pileipellis 92–155 µm wide, a trichodermium palisade composed of short, filamentous, septate, thin-walled, hyaline hyphae 3–5 µm in diam, arising from connective hyphae or inflated cells 14–29 µm in diam; terminal cells with bluntly acuminate, rounded or occasionally inflated apices, trichodermium becoming disorganized with age and somewhat gelatinized. Pileus context 70–110 µm wide, heteromerous, composed of interwoven, hyaline hyphae 2–3.5 µm in diam and abundant sphaerocysts 12–29 µm in diam in nests. Endocystidia absent. Stipitipellis composed of interwoven hyphae 2.5–4.5 µm in diam, terminal cells aggregated and projecting in places. Stipe-columella context heteromerous, of interwoven, hyaline hyphae 2–3 µm in diam and abundant sphaerocysts 17–25 µm in diam in nests. Hymenophoral trama 52–85 µm wide, heteromerous, of interwoven, hyaline hyphae 2–4 µm in diam and isolated nests of sphaerocysts 12–24 µm in diam; subhymenium well-developed, 18–28 µm wide, with 2–3 tiers of isodiametric cells 4–11 µm in diam (somewhat compressed). Clamp connections absent from all tissues. Basidia 21–39 x 8.5–12 µm, hyaline, clavate, with 4 sterigmata 4–7 µm long. Cystidia 60–87 x 10–17.5 µm, hyaline, broadly fusiform with acuminate or mucronate apices, sometimes clavate with or without mucronate apices, thin-walled, contents refractive in KOH; not or only slightly projecting beyond basidia, scattered. Spores 8–10.5 (12) x 6–7.5 (8.5) µm (9.1 ± 0.34 x 6.9 ± 0.22, n = 35), Q = 1.23–1.27, broadly ellipsoid, heterotropic, asymmetric, hyaline. Ornamentation amyloid, of warts 0.4–0.8 µm high, often 2 or more joined by fine, low lines to sometimes form a partial reticulum, with scattered isolated elements. Hilar appendix eccentric, conspicuous, tapered, 0.5–1 x 1–2 µm; plage indistinct, inamyloid. Spores cream in mass.

Etymology. Derived from the latin word ‘carmineus’, referring to the deep red color of the peridium.

Specimen examined: NEW ZEALAND. NELSON: Buller County, Karamea, Umere, epigeous, solitary or in small groups under Nothofagus menziesii, 8 January 1968, R.F.R. McNabb, PDD 26560 (HOLOTYPE: PDD).

Commentary. This was the first Macowanites to be described from Australasia. Macroscopically M. carmineus is much like a Russula, in its large basidioma, expanded pileus exposing the sublamellate gleba, and heterotropic spores. However, a spore print was unobtainable, and anastomoses between lamellae are common and formed irregularly. No other currently described species are this large, with as robust a stipe, lamellate to sublamellate gleba, and bright red peridium.

Macowanites carmineus resembles M. luteiroseus from Western Australia macroscopically, though it usually has a more robust pileus and stipe. Microscopically these two species differ in several features. Macowanites carmineus has a single-layered versus a two-layered pileipellis and less robust spore ornamentation with fewer connections between elements than M. luteiroseus.

Macowanites luteiroseus Bougher, Mycotaxon 63:37–48. Figs 1–6 , 1997. Figs. 25 , 26 .



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  		 FIG. 25. Macowanites luteiroseus A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 26. Scanning electron micrographs of spores of Macowanites luteiroseus. Bars = 10 µm.

 
Etymology. Derived from the latin words ‘luteus' and ‘roseus,’ meaning ‘yellow’ and ‘red’, referring to the color of the peridium.

Specimen examined: AUSTRALIA. WESTERN AUSTRALIA: Walpole-Nornalup National Park, Nunn Rd., emergent, in small or large groups under the leaf litter of wet eucalypt forests, 3 June 1995, K. Syme, PERTH 04259661 (HOLOTYPE: PERTH).

Commentary. A full description of the type of Macowanites luteiroseus can be found in Bougher (1997)Citation . Macowanites luteiroseus is a common and abundant fungus of the Eucalyptus diversicolor, E. guilfoylei and E. jacksonii forests of Western Australia. The overall appearance is generally that of an aborted Russula, however a spore print is not obtainable and the lamellae are much contorted and may have many connections between them. No other species has the combination of cream to yellow peridium with rosaceous patches, sublamellate gleba, abundant large hymenial cystidia, a two-layered pileipellis, and spores ornamented with isolated warts and partial reticulum.

Octaviania redolens G. Cunn., New Zealand J. Sci. Technol. 23B:172. pl 172b, 1942.

{equiv} Gymnomyces redolens (G. Cunn.) Pfister, Occ. Pap. Farlow Herbarium 9:43 1976.

{equiv} Martellia redolens (G. Cunn.) G.W. Beaton, Pegler and T.W.K.Young, Kew Bull. 39(4):682 1984.

= Stephanospora redolens (G. Cunn.) E. Horak, in Oberwinkler and Horak Pl. Syst. Evol. 131:163. 1979 comb. illegit. Figs. 27 , 28 .



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  		 FIG. 27. Gymnomyces redolens A. Basidioma. B. Peridiopellis and peridial context. C. Hymenophoral trama and hymenium. D. Basidia. E. Hymenophoral cystidia. F. Spores. Bars: A = 10 mm; B–F = 10 µm

 


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  		 FIG. 28. Scanning electron micrographs of spores of Gymnomyces redolens Bars = 10 µm.

 
Basidiomata 5–25 mm in diam, subglobose or irregular, depressed around basal attachment, rarely with an exposed gleba. Peridial surface slightly tomentose, finely wrinkled or smooth, white to cream, drying pale ochraceous. Context thin, fragile, off-white. Gleba white to cream becoming pallid ochraceous, loculate, locules small, irregular. Stipe absent. Columella absent or present, rudimentary, percurrent. Odor pleasant, resembling dried apricots; taste not recorded. Latex absent. Peridiopellis 70–150 µm wide, a dense turf of upright to repent, hyaline hyphal tips 20–55 x 2–4.5 µm in diam, becoming tangled and interwoven. Peridial context 210–450 µm wide, of tightly interwoven, nongelatinized, hyaline hyphae 2–3.5 µm in diam, sphaerocysts 12–22 µm in diam in scattered nests. Endocystidia absent. Columella context when present of interwoven, hyaline hyphae 2–3 µm in diam. Hymenophoral trama 40–75 µm wide, of interwoven, hyaline hyphae 2–4.5 µm in diam, nongelatinized, with sphaerocysts 12–27 µm in diam in scattered nests; subhymenium poorly developed, 11–20 µm wide, with 1–2 tiers of isodiametric cells 5–9 µm in diam. Clamp connections absent from all tissues. Basidia 25–35 x 9–11 µm, hyaline, ventricose to clavate, mostly with 1 or rarely 2 robust sterigmata 3–5 x 1–2 µm. Cystidia 24–49 x 8–15 µm, broadly ventricose with broadly rounded apices and few granular contents refractive in KOH; arising in subhymenium, not extending much beyond basidia, rare. Spores 9–12.5 x 9–12 µm (10.7 ± 0.3 x 9.9 ± 0.3, n = 35), Q = 1.05–1.07, globose to subglobose, orthotropic, symmetric, hyaline. Ornamentation amyloid, a dense spiny reticulum of warts and spines 2–4 µm high, bases nearly coalescing or several joined at their bases by low lines 0.2–0.8 µm high or ridges ±1–1.5 µm high in a partial reticulum. Hilar appendix 1–1.5 x 0.5–1 µm, cylindrical; plage inamyloid. Spores hyaline in mass, appearing pale cream when dried.

Etymology. Derived from the latin term ‘redolens,’ meaning ‘diffusing an odor.’

Specimens examined: NEW ZEALAND. AUCKLAND: Specimens collected among debris on the forest floor adjoining tracks in rainforest at the base of Mt Te Aroha (350 ft), May 1940, G. Cunningham 10141 (HOLOTYPE: K; ISOTYPE: PDD).

Commentary. Pfister (1976)Citation transferred this species to Gymnomyces, because G. redolens has amyloid spore ornamentation and sphaerocysts in the hymenophoral trama, both of which characters Octaviania lacks. Beaton et al (1984)Citation transferred it to Martellia, as they could find no sphaerocysts. However, the description by Beaton et al (1984)Citation is based on two collections (Beaton 27 and Beaton 31) from Victoria that do not match the holotype of Octaviania redolens G.Cunn. These two collections have an epithelial peridiopellis rather than a turf, more isolated spore ornamentation and more globose spores, 4- rather than 1- or 2-sterigmate basidia, and larger, more abundant cystidia than G. redolens. All these features are characteristic of the genus Cystangium rather than Gymnomyces. The recombination by Beaton et al (1984)Citation is validly published, but is not based on the type of O. redolens. The transfer by Pfister to Gymnomyces is considered to be a better placement of this species.

The recombination by Horak in Oberwinkler and Horak (1979)Citation , in the genus Stephanospora is illegitimate because full citation of the basionym is missing. Taxonomically G. redolens does not belong in Stephanosporaceae because of the presence of amyloid spore ornamentation characteristic of the Russulales.

The isotype examined from Auckland herbarium (PDD) matches the holotype in every respect. Gymnomyces redolens may be distinguished from other described species of Gymnomyces by the pallid ochraceous gleba at maturity, dense spiny reticulate spore ornamentation, basidia with 1–2 sterigmata and the association with Nothofagus