Further sporocarpic species of Glomus (Endogonaceae) from South Australia

[ 123 ] Trans . Br. mycol. Soc. 87 (1), 123-129 (1986)

Primed in Grear Br itain

FURTHER SPOROCARPIC SPECIES OF GLOMUS (ENDOGONACEAE) FROM SOUTH AUSTRALIA By P. A. McGEE Department of Plant Pathology, Waite Agricultural R esearch Institute, University of Adelaide, Glen Osmond, South Australia 5064

Three new species of Glomus (Endogonaceae) which form sporocarps, G. arborense, G. cerebriforme and G. warcupii, are described. The description of G. tenerum is emended and details are given of other South Australian species of Glomus which form sporocarps. A dichotomous key is included. Tandy (1975) described two new species of Glomus which formed sporocarps; G. tenerum Tandy (as tener) and G. tubiforme Tandy (as tubaeformis). She also described southern Australian collections of G. macrocarpum Tul. & Tul. sensu lato which Berch & Fortin (1983) later identified as G . caledonium (N icol. & Gerd.) Trappe & Gerd. Trappe (1982) omitted both G. tubiforme and G. tenerum from his synoptic key on the grounds that the type material did not match the published description. Trappe (pers. comm.) also believed that the holotype of G. tenerum (AD W 16428 ) was immature, possibly being an aberrant form of G. puloinatum (Henn.) Trappe & Gerd. Warcup (1985) considered biological and systematic details of G. tubiform e, and this species will not be discussed further. During 1982, 1983 and 1984 sporocarps were collected and pot cultures attempted using Metaleuca uncinata R.Br. (Myrtaceae) as the host plant. Isolates from various trap plants were also established in pot culture and checked for sporocarps up to 12 months after establishment. All soils and sporocarps came from collections in the Adela ide Hills, east of Adela ide, South Australia. The sporocarp-forming species of Glomus which were collected or recovered were G. arborense sp.nov., G. caledonium, G. cerebriforme sp.nov., G. clarum Nicol. & Schenck, G. macrocarpum, G. microcarpum Tul. & Tul., G. tenerum and G. warcupii sp.nov.

Glomus arborense sp.nov,

(Figs 1-3)

Sporocarpium hypogaeum ad 2 x 1 X 1 em, hyphae laxae aut confertae, radices et solum et sporas includentes, in ollis crescentes. Chlamydosporae terminales aut subterrninales, hyalinae aut luteolae, globosae aut subglobosae 26 x 24-55 x 50 !tm. Spora e laeves, parum asperae, ad 211m crassae. Contentum hyalinum guttis globosis olei interdum segregata septa 5-2o /lm in hypha sub spora. Hyphae subtentae 2'5-4!tm diam , cylindricae leviter turgidae prope murum sporae singularis, muris hyalinis aut luteolis ad 111m crassis. Hyphae in solo 3-8!tm diarn, muris hyalinis aut luteolis ad 1 11m crassis.

Holotypus ex solo apud Kuitpoensern sylvam in Australia meridionali per P. McGee lecto, Feb. 1982, ADW 16916.

Hypogeous sporocarps to 2 X 1 X 1 em, of loose to tightly formed hyphal masses enclosing roots, soil debris and randomly distributed spores, are formed in pot culture. Chlamydospores form terminally or subterminally, are hyaline to pale yellow, globose to subglobose, 26 x 24-55 x 50 pst: diam. The single spore wall has a smooth to dull roughened surface, to 3 pm thick. Contents are hyaline with globular oil drops and are cut off infrequently by a septum 5-20 pm down the subtending hypha. Sub tending hyphae 2 ' 5-4 pm diam, cylindrical with a slight flare at the spore wall , with a single attachment per spore. The wall of the subtending hypha is hyaline to pale yellow, to 1 pm thick. H yphae in the soil 3-8 pm diam, with walls hyaline to pale yellow, to 1 pm thick. The fungus forms vesicular-arbuscular rnycorrhizae (V AM ) on M. uncinata, Trifolium subterraneum L. , Calocephalus brownii (Cass.) F. Muell. and Wahlenbergia sp. Vesicles which form in the root are thin-walled and vary in shape from globose to irregular, with long sausage-shaped terminal structures common (Fig. 3). Spores have only been found in pot culture. Differs from G. pallidum Hall (1977 ) in having an undifferentiated hypogeous sporocarp, thinner spore walls, finer subtending hyphae and more frequent septa (T able 1). Isolated to pot culture from the roots of a trap plant of Ca/ocephalus brownii grown in soil collected from Kuitpo Forest, near Adelaide, South Australia.

Glomus cerebriforme sp.nov.

(F igs 4--6)

Sporae in sporocarpio epigaeo in agro aut in cultura in laxa hypogea massa hypharum. Sporocarpae epigeae t- 5 em latae, strato hypharum firmo multilobato plicate 1-2 mm

Glomus spp. from S. Australia

124

Table 1 . Characteristics of some small-spored species of Glomus which form sporocarps Sporocarp size (max. em), placement, shape and colour G. cerebriforme

G. arborense

G. pallidum (Hall, 1977) G. microcarpum (Berch & Fortin, 1984) G, puluinatum (T rappe, 1982) G. diaphanum (Mort on & Walker, 1984)

1"5 Epigeous Infolded layer White to grey 2 Hypogeous Hyphal mass Soil colour

Spore size Peridium Absent

Absent

Present 2'5 Epigeous Lobed to irregular White 2 Present Epigeous White to brown 1 Epigeous Rounded White ?

Spore wall number, max. thickness (pm) Subtending hyphae Septum diarn (fl m ) and type and type 2 Pre sent 25-80 3-6 Globose to Outer < 4, laminate Persistent irregular Inner < 1, membranous Hyaline Infrequent 2'5-4 25-55 Globo se to 2, laminate Persistent subglobose Hyaline to yellow 1 Infrequent 5- 15 28-78 Globose 1-8, laminate Persistent Hyaline (p m ),

Present

shape and colour

20-75 Globose Hyaline to brown 50-125 Globose Hyaline

1 1-12, laminate

Absent

3-12 Persistent

1 3-8

Present

9-18

39-121 Globose

2 Outer 2-6"5, laminate Inner < 1, membranous

Present Inverted into sub tending hyphae

5-11 Not persistent

Table 2. Characteristics of some large-spored species of Glomus which f orm sporocarps Spore size (pm)

G. warcupii

200320

G. macrocarpum (Berch & Fortin, 1983) G. caledonium (Gerdemann & Trappe, 1974) G. australe (Berch & Fortin, 1983) G. oersiforme (Berch & Fortin, 1983) G. tenebrosum (Berch & Fortin, 1983) G. alarum (N icolson & Schenck, 1979)

70225 120390

Spore wall details and thickness (pm) Outer mucilaginous layer Outer hyaline wall 15-60, inner brown wall 3-15 Outer wall hyaline thin Inner wall yellow 6-12 laminated Outer hyaline wall 1-8 Inner wall brown 4-10

120- Outer hyaline wall 4-8 33° Inner brown wall 7-35 60- Outer hyaline wall 1-5 200

Inner wall yellow 5-7

200- Single laminated yellow270 brown wall 13-z6

Spore closure Septum Brown

Spore Sporocarp arrangement development in sporocarp Aerogenous Random Ep igeous

Subtending hyphae Lateral short hyphae Present, cylindrical

7flm

Occluded

? Epigeous

Random

Random Septum ? Yellow Epigeous to 4flm Occluded Aerogenous? Clustered Epigeous Septum? Hyaline Thin Open

70- Outer mucilaginous layer Septum Hyaline 290 Outer hyaline wall 5-20 Thin Inner yellow wall 2-9

Cylindrical Cylindrical Lateral short hyphae present

? Epigeous?

Random

Cylindrical

? Epigeous?

Random

Cylindrical

Random Hypogeous

Random

Cylindrical

P. A. McGee crasso, centro cavo. Plexus hypharum inter solum et sporocarpium, sine peridio, album aut canum, solum includens. Chlamydosporae 1-8 in hyphis racemosis, confertae per sporocarpium, plerumque globosae, nonnullae irregulares 25 x 25-65 x 80 11m diam. Murus exterior sporae 2-4 11m crassus, hyalinus laminatus, laevis et in rnaturitate sordidus et asper. Murus interior sporae hyalinus membranaceus contentum viscidum et crystallinum in maturitate capiens. In hyphis subtentis collum sub septum membranaceum 0-811m sub apice. Hyphae subtentae 3-711m diam, 30-200 11m longae, hyalinae, cylindricae leviter turgidae prope murum sporae. Hyphae interdum infiatae et irregulares. Murus hyphae 211m crassus, sed 50 11m sub apice ad minus quam 1 11m decrescens. Holotypus sub Eucalypto sp. apud Kuitpoensem sylvam in Australia meridionali per J. H. Warcup, July 1984, ADW 16898.

Spores are found in epigeous sporocarps in the field or in loose hypogeous hyphal masses in pot culture. Epigeous sporocarps are to 1'5 em wide and have a firm, much lobed and infolded layer, 1-2 mm thick, around a hollow central area, lacking peridium. A plexus of hyphae connects the soil and the sporocarp layer. The sporocarp is white to dirty grey green colour and incorporates soil particles. Chlamydospores, found on racemose hyphae with one to eight spores, are packed tightly throughout the sporocarp layer. Spores mostly globose, some irregular, 25 x 25-65 x 80 pm diam. The outer spore wall is 2-4 pm thick, hyaline, laminate, smooth becoming dull and roughened with age. A hyaline, membranous inner wall cuts off the contents which become viscid and crystalline with maturity. A collar forms inside the subtending hypha supporting the membranous septum but may be up to 8 pm down the subtending hypha. Subtending hyphae are 3-7 pm diam, 30 to more than 200 pm long, hyaline, cylindrical with a slight flare at the spore wall, though occasionally swollen to irregular in shape, with a wall to 2 pm thick which thins to less than 1 pm thick within 50 pm of the spore. Specimens examined: Holotype: under Eucalyptus, Kuitpo, July 1984, J. H. Warcup (ADW 16899); under Eucalyptus, Mt Lofty, July 1984, P. McGee (ADW 16900); under Eucalyptus, Mylor, Aug. 1984, P. McGee (ADW 16901); under Eucalyptus, Cherryville, Aug. 1984, P. McGee (ADW 16902); under Eucalyptus, Piccadilly, Aug. 1984, P. McGee (ADW 169°3); under Eucalyptus, Greenhill, Aug. 1984, P. McGee (ADW 169°4).

The fungus forms VAM with M. uncinata, T. subterraneum and Wahlenbergia sp, In pot culture, external vesicles are globose, hyaline, thin-walled 25-45 pm diam, with external hyphae 1'5-7 pm diam. Penetration of the root can be either intercellular or direct into the epidermal cells via a swollen (to 7 pm) hypha to form hyphal loops.

125

Vesicles in the root are rare, never clustered, globose to subglobose, subterminal, thin-walled, 30 x 30-50 x 35 pm. The outer spore and hyphal wall of this fungus swell in lactic acid and occasionally an outer flaky layer is apparent. All field collections of sporocarps have been made in areas burnt in the bush fires of February 1983. Up to 550 sporocarps over 1 mm in diam per square metre have been counted in the field, 18 months after the fires.

Glomus warcupii sp.nov,

(Figs 7, 8)

Sporocarpium epigaeum rotundaturn ad 8 mm diam, 4 mm crassum, perideo gossypino tenui. Sporae in albida matrice hypharum fortuito dispositae, acrogenae, in maturitate 200-320 11m diam, gIobosae aut subglobosae, strato persistente exteriore hyalino mucilagino ad 15 11m crasso lamelloso. Primum murus sporae exterior lamellatus hyalinus 15-35 (60) 11m crassus in hypham subtentam descendens. Deinde in maturitate murus sporae interior lamellatus fuscus 3-15 pm crassus, crassissimus ad juncturam hyphae, usque ad 100 11m descendens, flavescens, tenuis. Contentum sporae hyalinum abscissum septo fusco lamelloso utrinque concavo ad 7 11m erasso, solum in sporis maturis, Hyphae subtentae cylindricae aut turgidae 20-50 pm diam, muro exteriore hyalino ad 12 pm crasso, muro interiore fusco aut luteolo 2-12 11m crasso, lumine 18-25 11m diam. Tenues rami ad 311m diam et 20 11m longi raro in hyphis subtentis, In sporocarpio hyphae hyalinae lamella tenui exteriore mucilagina, muro exteriore ad 5 11m crasso et muro interiore membranaceo, lumine 8-17 11m diam. In solo hyphae hyalinae 4-7 11m diam, duobus muris hyphae in toto 1 11m crassis. Vesiculae subterminales globosae muris tenuibus 25-4011m diam in hyphis cylindricis 3'5-5I1m diam. Vesiculae in radicibus raro 25-40 pm diam, globosae aut ovoideae. Ubique lamella exterior mucilagina. Holotypus sub Eucalypto sp. apud Kuitpoensem sylvam in Australia meridionali per J. H. Warcup, ADW 16905.

Epigeous rounded sporocarps to 8 mm diam and 4 mm thick, with a thin cottony peridium. Spores are randomly arranged in a white hyphal matrix and develop acrogenously, terminal on hyphae. Mature spores 200-320 pm diam, globose to subglobose with an ever-present, outer hyaline mucilaginous layer to 15 psx: thick which becomes flaky with maturity. The outer laminated hyaline spore wall develops first, 15-36 (60) pm thick and extends down the subtending hyphae. The laminated brown inner spore wall develops as the outer wall reaches maximum size, 3-15 pm thick, being thickest at the junction with the attached hypha, and extends up to 100 pm down the attached hypha, becoming pale yellow and thin. Contents of spores hyaline and cut off by a brown laminated septum, up to 7 pm thick, which is concave on both sides. Sub tending hyphae cylindrical to swollen, 20-50 pm diam, with a hyaline outer wall to 12 pm thick and

126

Glomus spp. from S. Australia

2

4

3

7

9

5

P. A. McGee

127

a brown to pale yellow inner wall 2-12 pm thick with a lumen 18-25 flm diam. Fine hyphal branches up to 3 flm diam by 20 flm long are infrequently observed on subtending hyphae. Hyphae in the sporocarp hyaline with a thin outer mucilaginous layer, an outer wall to 5 flm thick and a membranous inner wall with a lumen 8-17 flm diam. In soil, hyphae are hyaline 4-7 flm diam, with two hyphal walls a total of 1 flm thick. Subterminal globose thin-walled vesicles, 25-40 flm diam, form on cylindrical hyphae of 3' 5-5 flm diam in the soil. Vesicles 25-40 flm diam, globose to ovoid, form infrequently in roots. All structures have a mucilaginous outer layer.

Sporocarps up t04'5 x 2'5 x 1 cm(Fig. 9) are found most commonly some 18 months after summer fires. Soil particles and loosened, compressed spores at various stages of maturity are randomly arranged throughout a thin-walled hyphal matrix. Spores yellow, orange to brown, globose when distended, rarely pyriform, (45) 100-15° (200) flm diam. Spore contents hyaline, densely granular and become viscid with maturity and cut off by a plug (Fig. to). Subtending hyphae are cylindrical, 4-8 flm diam, with yellow/orange walls 1-2 flm thick at the spore wall, thinning to less than 1 flm. Globose thick-walled vesicles, 35 x 45-55 x 75 flm diam, form in roots of T. subterraneum.

Specimens examined: Holotype: under Eucalyptus, Kuitpo, July 1984, J. H. Warcup (ADW 16906); under Eucalyptus, Mt Lofty, Sept. 1984, P. McGee (ADW 16907); under Eucalyptus, Ironbank, Oct. 1984, P. McGee (ADW 16908); under Eucalyptus, Ironbank, Oct. 1984, P. McGee (ADW 16909).

Specimens examined: Lectotype; under Eucalyptus, Greenhill, Aug. 1984, P. McGee (ADW 16910); under Eucalyptus, Mylor, Aug. 1984, P. McGee (ADW 16911); under Eucalyptus, Kuitpo, J. W. Warcup (ADW 16912); under Eucalyptus, Cherryville, Aug. 1984, P. McGee (ADW 16913); under Eucalyptus, Piccadilly, Aug. 1984, P. McGee (ADW 16914).

The fungus forms VAM on a wide range of hosts including M. uncinata, T. subterraneum and Wahlenbergia sp. Of the large-spored sporocarpic species of Glomus, G. warcupii is distinct (Table 2) in having an outer mucilaginous layer over all structures, a thicker outer spore wall than inner spore wall, aerogenous spore development in sporocarps, a thick (to 7 flm) septum and lateral short hyphae adjacent to the spore on the subtending hyphae. Ectocarpic spores have not been observed. GLOMUS TENERUM Tandy, Aust. Jl Bot. 23: 864 (Figs 9, 10) (1975), emend. McGee Sporocarps found on the soil surface as rounded structures, young forms have a white cottony peridium, to 2 mm thick, of hyaline thin-walled hyphae, 5-7 flm diam, the hyphae becoming flattened and yellow with maturity. As the sporocarp grows the peridium breaks apart and is eventually only found as traces at the soil surface.

The holotype of G. tenerum (ADW 16428) has deteriorated in storage such that only a single spore from the dried material remains. The spore is now mounted on a slide in PV A lactophenol. The dried material of the paratype (ADW 16435) is extant though the portion stored in F .A.A. has also deteriorated. As one spore does not give sufficient information, the paratype becomes the specimen on which the species should be based. However, ADW 16435 varies from the type description in several ways. Further collections of G. tenerum have been made and are similar to ADW 16435. The description of G. tenerum is therefore emended, based on the paratype ADW 16435 which is thus selected as lectotype. The fungus forms VAM with a range of hosts including M. uncinata, T. subterraneum and Wahlenbergia sp. Epigeous sporocarps have formed in pot culture after two months, though ectocarpic spores have not been observed. Glomus tenerum differs from G. pulvinatum in

Fig. 1. Mature spore of Glomus arborense, subterminal on degenerate hypha (x 1370). Fig. 2. Septum of the spore of Glomus arborense in Fig. 1 (x 1370). Fig. 3. Terminal hyphal swelling of Glomus arborense in a cortical cell of Trifolium subterraneum (x 1000). Fig. 4. Squashed spore of Glomus cerebriforme showing the thicker outer wall, membranous inner wall (arrowed) and viscid contents of a mature spore (x 1000). Fig. 5. Septum (arrowed) and supporting collar of a mature spore of Glomus cerebriforme (x 1000). Fig. 6. Hyphae of Glomus cerebriforme mounted in lactic acid showing racemose spore development and swelling of the outer spore wall and hyphal wall ( x 400). Fig. 7. Spore of Glomus warcupii showing the dark inner spore wall, hyaline outer spore wall with mucilaginous layer (arrowed), septum and short, lateral projection which is often found on the subtending hypha (x 400). Fig. 8. Echinulations projecting into the spore contents of Glomus warcupii (x 1000). Fig. 9. Sporocarp of Glomus tenerum showing the open, amorphous nature of the mature sporocarp ( x r), Fig. 10. Details of the plug (arrowed) and spore wall of Glomus tenerum ( x 1000).

128

Glomus spp. from S. Australia

having two walls, larger spores, a plug cutting off spore contents and finer-subtending hyphae. Further species of Glomus which form sporocarps have been collected in South Australia and include the following.

restricted by wall expansion. Suhtending hyphae are hyaline to yellow, 6-12 pm diam with walls to 2 pm thick.

GLOMUS CALEDONIUM (Nicol. & Gerd.) Trappe & Gerd., Mycol. Mem. S: 56 (1974). Sporocarps are rounded, up to 4 mm diam, with a white cottony exterior with spores randomly distributed in a white cottony interior. Spores are globose to subglobose, 15<>-220 pm diam, with a hyaline outer wall 2-5 pm, easily separated from a yellow to brown inner wall 3-5 pm thick. The outer wall extends down the subtending hypha 5<>-70 pm. No septum is apparent in fresh material. The contents are under great pressure in the spore and burst from severed hyphae. The attached hyphae are cylindrical, 25-35 pm diam. Attempts to pot-culture this fungus have failed.

The fungus was isolated to pot culture from a trap plant of Leptospermum sp., grown in soil collected from Lobethal by Dr J. H. Warcup.

Specimens examined: under Eucalyptus, Ironbank, 1969, ADW 16417; under Eucalyptus, lronbank, 1969, ADW 16418; under Eucalyptus, Loftia Park, 1969, ADW 16419 ; under Eucalyptus, Belair, 1971, ADW 16421; under Eucalyptus, Mt Crawford, 1971, ADW 16423; under Eucalyptus, Ironbank, 1974, ADW 16437.

GLOMUS CLARUM Nicol. & Schenck, Mycol. 71: 182 (1979)· Sporocarps of loose aggregations of hyphae enclosing soil particles and randomly arranged spores, up to 3 em long, and are formed hypogeously in pot culture. Chlamydospores formed in sporocarps or roots, are globose to subglobose to ovoid, 75-160 pm diam. The outer wall is 4-8 pm thick, hyaline to pale yellow, extends up to 10 pm down the subtending hypha and is covered by a mucilaginous layer to 4 pm thick. The yellow to brown inner wall is 4-8 pm thick. Contents are

Specimen examined: from soil collectedfrom Lobethal, 1982, J. H. Warcup, ADW 16917.

GLOMUS MACROCARPUM Tul. & Tul., Giorn. Bot. Ital. 2: 63 (1845). Sporocarp an open fragile structure, epigeous, without peridium, incorporating soil particles to 3 mm diam, with hyaline to pale yellow hyphae and brown spores. Chlamydospores globose to ellipsoid, 150 x 235-230 x 235 pm, with hyaline contents. The outer wall is hyaline to pale yellow, smooth to dull roughened, 1-2 pm thick, and is easily separated from inner wall, which is laminated, brown, 12-25 pm thick, being thickest at the base and occluding the contents. The inner wall extends down the subtending hypha as much as 100 uxt». Subtending hyphae are 2<>-25 pm diam. Attempts to pot-culture this fungus have failed. Specimen examined: under Eucalyptus, Woodside, 1983, P. McGee, ADW 16918.

GLOMUS MICROCARPUM Tul. & Tul., Giorn. Bot. Ital. 2: 63 (1845). Sporocarps of loose aggregations of hyphae enclosing soil particles and randomly arranged spores, up to 2 em diam, are formed hypogeously in pot culture. Spores vary from the type (Berch & Fortin, 1984) in that the spore wall is a maximum of 3 pm thick, without laminations and the subtending hyphae are 3-5 pm diam, with walls to 1 pm thick. Vesicles observed in roots can be from 25 x 25 pm to 50 x 85 us»,

KEY TO THE SPOROCARPIC SPECIES OF GLOMUS OF SOUTH AUSTRALIA Spores in sporocarp mostly 10-80 pm diam Spores in sporocarp mostly 70-350 p;m diam 2. Spores mostly with septum when mature 2. Spores without septum 3. Spores with one wall 3. Spores with more than one wall 4. Forms ectomycorrhizae, spores 10-40 pm diam 4· Forms VA mycorrhizae, spores 20-60 p;m diam 5. Mucilage over all structures 5. No mucilage 6. Forms epigeous sporocarps, spores in sporocarps mostly 2()(>-300 pm 6. Forms hypogeous sporocarps, spores in sporocarps mostly 80-150 pm 7· Subtending hyphae 15-35 p;m . 7· Subtending hyphae 4--9 p;m, spores yellowto orange 8. Sporocarp firm, rounded, with peridium, spores with fine outer wall, thicker inner wall. 8. Sporocarp fragile, open, lacking peridium, spores with two wallsof similar thickness 1.

2

1.

5 3 4

. G. arborense G. cerebriforme . G. tubiforme G. microcarpum · 6

· 7 G. warcupii G. clarum · 8 G. tenerum G. caledonium G. macrocarpum

P. A. McGee Specimen examined: from soil collected from Lobethal, 1982, J. H. Warcup, ADW 16919.

The fungus was isolated to pot culture from a trap plant of Leptospermum sp. grown in soil collected from Lobethal by Dr J. H. Warcup. I wish to thank Dr I.' R. Hall for providing figures 1 and 2, Mrs M. Marlow for providing the Latin translations and Mr D. Symon and Dr J. H. Warcup for reviewing the manuscript. REFERENCES

BERCH, S. M. & FORTIN, J. A. (1983). Leetotypification of Glomus macrocarpum and proposal of new combinations: Glomus australe, Glomus versiforme, and Glomus tenebrosum (Endogonaceae). Canadian Journal of Botany 61, 2608-2617. BERCH, S. M. & FORTIN, J. A. (1984). A lectotype for Glomus microcarpum (Endogonaceae, Zygomycetes). Mycologia 76, 190-193.

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GERDEMANN, J. W. & TRAPPE, J. M. (1974). The Endogonaceae of the Pacific Northwest. Mycologia Memoir S, 1-76. HALL,1. R. (1977). Species and mycorrhizal infections of New Zealand Endogonaceae. Transactions ofthe British Mycological Society 68, 341-356. MORTON, J. B. & WALKER, C. (1984). Glomus diaphanum: a new species in the Endogonaceae common in West Virginia. Mycotaxon Z1, 431-440. NICOLSON, T. H. & SCHENCK, N. C. (1979). Endogonaceous mycorrhizal endophytes in Florida. Mycologia 71,178-198. TANDY, P. A. (1975). Sporocarpic species of Endogonaceae in Australia. Australian Journal of Botany 23, 84
(Received for publication 17 October 1985)

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