Mycol. Res.
96 (3), 233-236 (1992)
233
Printed in Great Britain
Spiral-spored Tulasnella species from Devon and the New Forest
PETER ROBERTS 36 Western Road, Torquay, Devon TQl 4RL, U.K.
Tulasnella interrogans, T. anguifera and T. falcifera are described as new. T. helicospora is described and recorded as new to Britain.
Extensive colleding of Tulasnella Schroter species in South Devon, England, over the last three years has led the author to reassess current species concepts within the genus, placing particular emphasis on hypha!' basidial and hymenial morphology as well as spore shape and size. This initial contribution records four easily distinguished spiral-spored species, three of which appear to be undescribed. The parameters for these species are precise, and colledions diverging substantially from the descriptions and illustrations should be treated with caution. If just a handful of sites in the south of England can produce four spiral-spored Tulasnella species, there must undoubtedly be many more elsewhere that still await discovery and description.
The Devon material compares well with the type description and spore drawings in Raunkiaer (1918), based on an 1893 Danish colledion on fallen beech leaves. The difference is that the basidia were described as arising diredly from the basal hyphae, which in the Devon colledion would only be true in
Tulasnella helicospora Raunkiaer, Bofanisk Tidsskri/f 36: 205 (1918). (Fig. 1)
Gloeofulasnella helicospora (Raunkiaer)' Christiansen, Danish Resupinate Fungi. 1. Ascomycetes and Heterobasidiomycetes. Dansk Botanisk Arkiv 19: 40 (1959).
Basidiomes barely visible, no more than a thin, effused, pruinose to felted whitish-grey patch on the underside of a fallen branch. Hyphae undamped,. 3'6-6'0 I-lm diam., thinwalled, forming a conventionally branching, compad but easily visible hymenium. Basidia comparatively large (by volume), up to 10'8 I-lm across, mostly widely clavate, but varying from subglobose to narrowly clavate. Sterigma fa normally four per basidium, comparatively large (by volume), up to 6'6 I-lm across, globose or broadly davate when young, becoming clavate to mitriform, then variously extending at the tip to 20'0 I-lm or more, normally remaining swollen above the base. Spores normally 'question-mark'-shaped, tightly inrolled and spiralled at the tip, ca 20'0-30'0 x 1'8-3'6 I-lm (when notionally uncoiled), replicating in mount, but germinating by tube in a damp print. A small minority of spores in a large print deviate from the question-mark shape, mostly by having a more open loop. Specimen examined: on fallen Fraxinus branch, Slapton Wood, South Devon, 25 Nov. 1990. P. Roberts 113, K.
lOj.l.m Fig. 1. Tulasnella helicospora: basidia; replicating spores from mount; spores from print (Roberts 113).
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P. Roberts areas of new growth. As the hymenium develops and matures, these initial basidia-bearing side branches continue to branch, forming new basidia in succession. Tulasnella helicospom is easily distinguished from other known spiral-spored species by the wide unclamped hyphae and the rather large distinctively shaped spores. It clearly belongs to a group of species allied to T. violea (Quel.) Bourd. & Galz. (interpreted pro tern. as a species with large globose spores), all of which share the same branching hymenial morphology, the same wide and easily visible unclamped hyphae, and the same voluminous basidia and sterigmata. Indeed, T. helicospom is microscopically indistinguishable from T. violea, except for the spores. Rogers (1933), followed by Olive (1957), placed T. helicospom in synonymy with T. calospom (Boud.) Juel, a name to which they assigned virtually any Tulasnella with spores over 15 IJm long regardless of other features. This deserves no further comment. Tulasnella calospom forma spirillifem Bourd. & Galz. has been placed in synonymy with T. helicospora by all subsequent authors, apparently because it was briefly described as having spiral spores. This in itself is now insufficient, and collections so labelled should be re-examined. The Siapton collection of T. helicospora appears to be the first record of its occurrence in Britain.
Tuiasnella interrogans P. Roberts, sp. nov.
(Fig. 2)
Etym.: from the Latin, interrogans = questioning, referring to the question-mark shape of the spores Basidiomata resupinata, inconspicua vel invisibilis Iicet observetur sine lente, ceraceo-ge1atinosa, violaceo tinda. Hyphae fibulatae, angustae, 1'8 \.1m latae. Basidia obovata vel subglobosa, stipitata. Sterigmata (2) 4, primum globosa, post demum fusoidea. Sporae spiralliferae. T. helicospora Raunk. proximae. sed minores. 12'0-30'0 x 1'2-1'8 1Jrn.
Ad ligna putrida, Great Plantation, Bovey Tracey. Devon, Anglia, 2 Feb. 1991, P. Roberts 136, holotypus, K.
Basidiomes resupinate, effused, inconspicuous or invisible to the unaided eye. When visible, subgelatinous to ceraceous when wet, becoming matt and finely felted when damp, and finally drying to invisibility. The fruit body is distinctly violaceous when wet, becoming somewhat greyer with a whitish pruina when damp. Hyphae conspicuously clamped at all septa, consistently thin, around 1'8 IJm wide, forming a conventionally branching and easily visible hymenium when mature. Basidia normally distinctly stalked, globose, ovoid or irregular, often showing hyphal continuation as weak but distinct outgrowths or projections, rather small (by volume), up to 6'6 IJm or so wide. Sterigmata normally 4, but frequently 2 per basidium, globose to clavate when young, becoming clavate to mitriform, then variously extending to 12'0 IJm or more, up to 4'8 IJm or so wide, normally broadest just above the base, occasionally dividing to become bi-apiculate. Spores normally 'question-mark shaped', often extending into an open loop, rather similar to the spores of T. helicospora, but consistently smaller, narrower and without the tightly inrolled spiral at the end, 12'0-30'0 x 1'2-1'8 IJm when notionally
10J.1m Fig. 2. Tulasnella interrogans: basidia-bearing branching hyphae; replicating spore from mount; spores from print (holotype specimen).
extended. A small minority of spores deviate from the pattern, normally by forming a more open loop. Replicating in mount, germinating by one or more tubes in damp print. Specimens examined: on bark of Picea log, together with an undamped allantoid-spored Tulasnella species, Great Plantation, Bovey Tracey, South Devon (SX 8275), 5 Mar. 1989, P. Roberts 97, K; same locality, on decorticated fallen Corylus branch, 1 Apr. 1989. P. Roberts 98, K; same locality. on decorticated fallen Salix branch, 2 Feb. 1991, P. Roberts 136, holotype. K; on fallen Picea branch, Bellever Forest. Dartrnoor, South Devon (SX 6377), 13 Apr. 1991, P. Roberts 200, K.
Tulasnella interrogans is distinguished from T. helicospom by its thin clamped hyphae and smaller distinctively shaped spores. The presence or absence of an easily visible fruit body is simply a matter of growth and maturity. The holo type collection, with visible fruit body, has an extensively branched hymenium producing basidia in succession. The second collection (Roberts 98), with no visible fruit body, is clearly immature and consists only of a few trailing basal hyphae producing single basidia on side branches, some of which are beginning to show signs of branching further. An allantoid-spored species which is microscopically identical to T. interrogans has been collected in Devon, and undoubtedly these form part of a wider group of allied species having the same hypha!' hymenial and basidial morphology.
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Spiral-spored Tulasnella species
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10 Jlm
10 Jlm Fig. 3. Tulasnella anguifera: basidia showing grossly swollen sterigmata; basal hyphae showing first stages of hymenial growth; replicating spores from mount; spores from print (holotype specimen).
Fig. 4. Tulasnella falcifera: basidia-bearing branching hyphae with undulating cystidia; spores from print (holotype specimen).
T. interrogans is obviously indifferent to substrate, and extensive collecting in Devon suggests that this is true for all Tulasnella species.
becoming grossly swollen and variously shaped, clavate, widely fusiform, or elliptical and often appearing waisted, frequently larger than the basidium, up to 6'6 ~m or so wide, variously extending to 14 ~m or longer. Spores spiralling in 3 dimensions, often appearing coiled or forming an open loop, 13'2-30'0 x 1'8-3'6 ~m when notionally extended. A small minority of spores show varying coiled or spiralled patterns.
Tulasnella anguifera P. Roberts, sp. nov. (Fig. 3) Etym.: from the Latin, anguiferus = bearing snakes, with reference to the serpentine shape of the spores
Basidiomata resupinata, inconspicua vel invisibilis licet observetur sine lente, plusminusve ceraceo, lilaceo-grisea tinda. Hyphae fibulatae, angustae, 1'8-3'6 ~m latae. Basidia stipitata, plusminusve obovata. Sterigmata obovata vel clavata, saepe multis partibus maiora quam basidiis. Sporae spiralliferae, serpentiformes, 13'2-30'0 x 1'8-3'6 ~m. Ad ligna putrida, Stover Park, South Devon, Anglia, 29 Dec. 1990, P. Roberts 109, holotypus, K. Basidiome resupinate, effused, inconspicuous or invisible to the unaided eye, when visible, ceraceous to matt felted, dull grey-violaceous. Hyphae clamped at all septa, thin in hymenium, 1'8 ~m diam.; basal hyphae wider, to 3'6 ~m, and with thicker refractive walls, forming a conventionally branching hymenium with the hyphae clearly visible. Basidia normally more or less stalked, though often appearing narrowly clavate, rather small, to 4'8 ~m or so wide, disproportionately small in comparison with the sterigmata. Sterigmata four per basidium, initially globose to clavate,
Specimens examined: on fallen branch, probably Prunus spinosa, Grley Common, South Devon (SX 8266), 27 Dec. 1990, P. Roberts 110, K; on decorticated Picea log, Stover Park, South Devon (SX 8375), 29 Dec. 1990, P. Roberts 109, holotype K. Tulasnella anguifera is distinguished from T. helicospora by its thin clamped hyphae and from both T. helicospora and T. interrogans by its distinctively coiled and spiralling spores, which may well prove to be diagnostic for the species. A minority of spores in a print will diverge from this shape in various ways. The branching pattern of hymenial growth is conventional, but the basidial morphology is exceptional and so far unique among Tulasnella species collected in Devon. In mature sporebearing basidia, the grossly swollen sterigmata are quite out of proportion with the comparatively small basidium, often equalling or substantially surpassing it in volume. The holotype collection contains the remains of Dacrymyces stillatus Nees: Fr. fruit bodies which have been invaded by the Tulasnella. These reveal early growth stages with particular
P. Roberts
236
clarity. The initial penetrating hyphae are rather wide, somewhat thick-walled, rarely septate, and maintain exceptionally straight lines for several millimetres into the deliquescent Dacrymyces. Basidia are formed at intervals directly on these basal hyphae, the basidial stalk then branching to initiate the build-up of the hymenium (see Fig. 3). It seems probable that this represents the typical growth pattern for most Tulasnella species.
Tulasnella falcifera P. Roberts, sp. nov.
(Fig. 4)
Etym. : from the Latin, faiciferus = bearing sickles, with reference to the typically sickle or question-mark shape of the spores Basidiomata resupinata,inconspicua vel invisibilis licet observetur sine lente. Hyphae non fibulatae, angustae, 1'2-1'8 ~m latae. Basidia obovata vel subglobosa, stipitata. Sterigmata primum globosa, post demum fusoidea. Cystidia elongata, sinuosa, 24'0-54'0 x 6'0 ~m. Sporae spiralliferae, T. interrogans proximae, sed minores, 11'4-16'8 x 1'8-3'0
~m.
Ad ligna putrida, New Forest, Hampshire, Anglia, 1 Apr. 1991, T. Laessoe 2309, holotypus, K.
Basidiomes resupinate, inconspicuous or invisible to the unaided eye, in type specimen growing on or with Scopuloides hydnoides (Cooke & Massee) Hjortst. & Ryv. Hyphae unclamped and consistently thin, around 1'2-1'8 ~m diam., branching, with many 'H' connexions, not or barely seen to form a hymenium. Occasional clamp connexions may be present on basal hyphae, but these not clearly seen. Basidia globose, ovoid, or irregular, often rather narrow and elongated; normally clearly stalked; often showing hyphal continuation (Accepled 1B October 1991)
as distinct papilla, outgrowths or projections, rather small (by volume), up to 4'8 ~m or so wide. Sterigmata globose when young, becoming clavate to mitriform, then variously extending to 12'0 11m or more, up to 3'6 11m or so wide, normally broadest just above the base. Cystidia elongated, sinuous or undulating, obtuse, 24'0-54'0 x 6'0 ~m; with distinct but weakly refractive contents. Spores normally 'question-mark' or sickle shaped, often with a tight threedimensional loop at the tip; more or less identical to the spores of T. interrogans; 11'4-16'8 x 1'8-3'0 11m when notionally extended, Tulasnella faicifera is easily distinguished from other spiralspored species by the presence of cystidia and its thin unclamped hyphae. The only other cystidiate species so far recorded for Britain is T. cystidiophora Hahn. & Litsch., which has subglobose spores. The spores of T. faicifera are remarkably similar to those of T. interrogans, though on average slightly smaller and with a greater tendency to form a final tightly spiralled loop. For all practical purposes they are indistinguishable in a print. Thanks to David Minter for suggestions and amendments to the Latin text and to Thomas Laessoe for forwarding interesting new material.
REFERENCES Bourdot, H. & Galzin, A. (I928). Hyminomyceles de France. Sceaux. Olive, L. S. (1957). Tulasnellaceae of Tahiti. A revision of the family. Mycologia 49, 663--679. Raunkiaer, C. (1918). En ny Tulasnella-Art samt Bemaerkninger om Tulasnella's systematiske Stilling. Bolanisk Tidsskrifl 36, 204-212. Rogers, D. P. (1933). A taxonomic review of the Tulasnellaceae. Annales Mycologici 31, 181-203.