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Allantoid-spored Tulasnella species from Devon
213
Mycol. Res. 97 (2): 213-220 (1993) Printed in Great Britain
Allantoid-spored Tulasnella species from Devon
PETER ROBERTS 36 Western Road, Torquay, Devon TQI 4RL
Tulasnella permacra, T. saveloides, and T. fomaculum are described as new. T. danica is described and recorded as new to Britain. T. allantospora is redescribed with reference to the holotype. Extra-limital species are briefly noted and a key is provided to accepted taxa.
Extensive collecting 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. More emphasis is placed on hyphal, spore, and hymenial morphology and less on fruit-body size, colour, texture, and habitat. An initial paper (Roberts, 1992) looked at four species with spiral spores. This second contribution examines five allantoidspored species found in Devon, with reference to collections made elsewhere in Britain plus published descriptions of overseas species. Tulasnella appears to be a genus rich in species and it is highly unlikely that a few woodlands in South Devon will have yielded up more than a small proportion of them. Care should therefore be taken before automatically assigning collections from elsewhere to the following species.
Tulasnella allantospora Wakef. & Pears., Trans. Br. Mycol. SOC.8 : 220 (1923) (Fig. I) Gloeotulasnella caroliniana Olive, Bull. Torrey Bof. Club 80 (1): 41 (1953).
Basidiomes often invisible to the naked eye, or forming no more than a faint, dull bloom. When visible, effused, ceraceous or subgelatinous, violaceous grey. Hyphae clamped at all septa, ca 2.0-3.0 pm diam., forming an extensively branched, basidia-bearing hymenium. Basidia variously clavate to subglobose, ca 4-0-6.5 vm wide, with or without a short or long stalk, clamped at the base. Sterigmata 4 per basidium, ellipsoid becoming fusiform, ca 3.0-4.0 pm diam. at widest point, variously extending up to 12.5 ym or more. Spores weakly allantoid, tapered at both ends, averaging 6.5-9.0(-10.0) x 2.5-3.0 pm (see Table I), replicating in mounts. Specimens examined: on fallen Pinus branch, Great Plantation, Bovey Tracey, South Devon (SX 8275), 12 Jan. 1991, P. Roberts 124, K; same location and substratum, 16 June 1991, P. Roberts 240, K; on decorticated Picea log with Tylospora jbrillosa (Burt) Donk and Tulasnella sp., Bellever Forest, Dartmoor, South Devon (SX 6376). 27
Apr. 1991, P. Roberts 220, K; same location, on fallen Picea branch, I Dec. 1991, P. Roberts 340, K; on fallen coniferous fence-post, Orley Common, South Devon (SX 8266), 9 Nov. 1991, P. Roberts 318, K; same location, on fallen deciduous branch, 30 Nov. 1991, P.Roberts 339, K ; on fallen branch, probably Picea, with Botyobasidium subcoronatum (Hohn. & Litsch.) Donk, Fernworthy Forest, Dartmoor, South Devon (SX 6583), 1 Mar. 1992, P. Roberts 365, K; on decorticated coniferous wood, East Horsley, Surrey, 4 Apr. 1922, A. Pearson, holotype, K.
Tulasnella alhnfospora has been a much-confused species ever since it was first described. Wakefield & Pearson (1923) based their type description on two specimens collected from the same locality a week apart. Unfortunately these represent two different species. As already pointed out by Rogers (1933), the earliest specimen (taken to be the holotype) possesses abundant and clearly visible clamp-connexions. The second collection, on which Wakefield & Pearson's description and illustration seem to be based, has similar spores but lacks clamp-connexions. As a result, T. allantospora has been variously interpreted in the literature and all records are suspect unless accompanied by a full description or voucher material. Based on the holotype, T. allantospora is a consistent and easily recognized species, distinguished by its conventionally branching hymenium with clamps at all septa, together with its weakly allantoid spores, the majority of which (in every print) taper towards each end, more particularly towards the apiculus. It has also proved to be one of the commonest Tulasnella species in Devon, and the collections cited above are just a representative sample of the total seen. T. interrogans P. Roberts is a closely related but less common species with spiral spores. Bourdot & Galzin (1923) originally described Tulasnella rubropallens Bourd. & Galz. as having similar spores (measuring 7.5-9 x 3-4 pm) but with only scattered clamp-connexions on the hyphae and none at all on the basidia. The hyphae and basidia were described as wider. Later (Bourdot & Galzin,
Allantoid-spored Tulasnella species from Devon
P-, A
Fig. 1. Tulasnefla allantospora. A, Basidium and spores from mount (holotype specimen); B, (compact fruit body) branching hyphae, basidia, and spores from print (P. Roberts 124); C, (more open fruit body) basidia, spores from print, and hyphae showing continued growth through the basidia (P. Roberts 318).
Table I . Tulasnella allantospora: spore measurements (pm) from prints of the collections cited plus relevant publications Holotype* P. Roberts 124 P. Roberts 220 P. Roberts 240 P. Roberts 318 P. Roberts 339 P. Roberts 340 P. Roberts 365 Jiilich (1984) Olive (1953)t * Measurements taken from a few scattered spores in a mount. The original published spore measurements almost certainly refer to a different, unclamped species t Published measurements for holotype of Gloeotulasnella caroliniana.
1928), they suggested their species might be conspecific with T. allantospora, a view shared by most subsequent authors. Jiilich (1984), however, has repositioned T. rubropallens as an entirely unclamped species with spores measuring 7-9(-10) x 2.4-3.4 pm, though on what basis is unclear. It urgently requires a modem redescription. Olive (1953) described an American species, Gloeofulasnella caroliniana Olive, which appears to have identical spores and is distinguishable from T. allanfospora only by its subgelatinous
fruit body. Since fruit-body texture has more to do with humidity than taxonomy, G. caroliniana is here proposed as a synonym of T. allantospora. Tulasnella permacra P. Roberts, sp. nov.
(Fig. 2)
Etym.: from the Latin, per plus macer = very meagre
Basidiomata resupinata, inconspicua vel invisibilia licet observetur sine lente. Hyphae fibulatae, angustae, 0.5-1.5 pm latae. Basidia stipitata, obovata vel clavata, angusta, ca 2.5-4.0 ym lata. Sferigmafa 4, fusoidea. Sporae allantoideae, T. allantospora proximae sed minores. ca 5.5-8.0 x 2.0-2'5 pm. Ad ligna ~utrida,Orley Common, Devon, Anglia, 19 Jan. 1991, P. Roberts 125, holotypus, K. Basidiomes resupinate, inconspicuous or invisible to the unaided eye. Hyphae clamped, unusually and consistently thin, ca 0.5-1.5 pm diam., sparsely branched, not seen to form a hymenium but trailing through and over the substratum, producing basidia individually on short side branches. Basidia narrowly clavate or ovoid, ca 2.5-4.0 pm wide, with a distinct stalk, clamped at the base. Basidia frequently appear elongated or narrowly fusiform as a result of apical hyphal outgrowths. Sterigmata 4 per basidium, ellipsoid, becoming widely fusiform, ca 3.0-4.5 pm diam. at the widest point, variously extending up to 8.5 pm. Spores similar to T. albntospora but generally shorter, narrower, and more strongly curved, ca
Peter Roberts
Fig. 2. Tulasnella permacra. A, Typically thin hyphae with ovoid and elongated basidia, plus spores from print (holotype specimen); B, basidia and spores from print (P. Roberts 252).
5.5-8.0 X 2.0-2.5 ym; replicating in mounts. (6-0-)6.58-0(-8.5) x (2-0-)2.5(-3-0) ym (P. Roberts 125, holotype); 5.5-8.0 x 2.0(-2.5) ym (P. Roberts 252); 5.0-6.0 x 2.02.5 ym (P. Roberts 312).
Specimens examined: on fallen decorticated Picea branch with Tulasnella violea (QuCl.) Bourd. & Galz., Stover Park, South Devon (SX 8375), 11 Nov. 1989, P. Roberts 101, K; on fallen Fraxinus branch, Orley Common, South Devon (SX 8266), 19 Jan. 1991, P. Roberts 125, holotype, K; same location, on fallen deciduous wood with Tulasnella sp. and Helicogloea lagerheimii Pat., 22 June 1991, P. Roberts 252, K; same location, on fallen coniferous fence post with Xenosperma ludibundtrm (Rogers & Liberta) Oberw. ex Julich and Iulasnella sp., 27 Oct. 1991, P. Roberts 312, K. Tulasnella pemacra is a cryptic but distinctive species, easily separated from T. allantospora b y its exceptionally thin, clamped hyphae, tiny and unusually narrow basidia, and rather smaller, narrower, and more strongly allantoid spores. The few collections made, all of which are cited above, consist solely of trailing, sparsely branched hyphae producing basidia singly o n short side branches. However, 7.permacra does not appear t o be an obligate hymenial parasite and there is n o intrinsic reason why it should not form a more compact hymenium when growth conditions permit. T h e exceptionally narrow basidia are a constant feature of the species. Their narrowness is exacerbated b y a tendency for hyphal growth t o continue through the basidium, emerging a t
Fig. 3. Tulasnella tomaculum. A, Basidia-bearing hyphae, replicating spore, basidia showing variable sterigmata, and spores from print (holotype specimen); B, young, open growth in the hymenium and spores from print (P. Roberts 341).
the apex (or occasionally laterally) as a papillate or snout-like outgrowth. Such basidial outgrowths are common in Tulasnelb species, and are particularly prevalent in young o r sparse collections, less evident or absent when a stable hymenium is formed. The formation of sterigmata usually prevents further hyphal growth from the basidium; where growth does continue, the basidium is normally left as a sterile cell or reduced t o little more than a swelling. Very occasionally immature sterigmata can be found o n such swellings (see T . allantospora, Fig. Ic), but none has yet been seen t o produce spores.
Tulasnella tomaculum P. Roberts, sp. nov.
(Fig. 3)
Etym.: from the Latin, fornaculum = a kind of small sausage, with reference t o the allantoid spores
Basidiomafa resupinata, inconspicua, ceraceo-gelatinosa, lilaceo-grisea tincta. Hyphae non fibulatae, ca 2.5-3.5 pm latae. Basidia clavata vel obovata, ca 4.0-5.5 pm lata. Sterigmata 4, fusoidea vel clavata. Sporae breviter allantoideae, ca 4.0-7.0 x 2.5-3.0 pm. Ad ligna putrida, Orley Common, South Devon, Anglia, 27 Dec. 1990, P. Roberts 119, holotypus, K. Basidiomes often inconspicuous o r invisible t o the naked eye. When visible, effused, resupinate, ceraceous t o subgelatinous,
216
Allantoid-spored Tulasnella species from Devon violaceous-grey. Hyphae unclamped, forming a conventionally branching and often compact basidia-bearing hymenium. Basal hyphae normally long and straight, somewhat thickwalled in mature specimens; subh~menial hyphae rather short-celled and slightly swollen, ca 2-5-3.5 pm diam. Basidia normally clavate, but frequently ovoid or subglobose, ca 4.0-5.5 pm wide, often with a poorly defined long or short stalk. Sferigmata 4 per basidium, variable but normally fusiform, sometimes elongated or clavate, ca 2.5-3.5 pm diam. at the widest point, variously extending- up. to 21.0 pm or more. Spores weakly allantoid, normally obtuse at each end, ca 4.0-7.0 x 2.5-3.0 pm (see Table 2), replicating in mounts. Specimens examined: on bark of Picea log, Great Plantation, Bovey Tracey, South Devon (SX 8275), 1Apr. 1989, P. Roberts 96, K; same location and substratum, 16 June 1991, P. Roberts 238 & 239, K; on fallen Picea branch, Stover Park, South Devon (SX 8375), 1Dec. 1990, P. Roberts 118, K; on decorticated deciduous branch, possibly C o y l w , Orley Common, South Devon (SX 8266), 27 Dec. 1990, P. Roberts 119, holotype, K; same location, on fallen deciduous wood, 30 Nov. 1991, P. Roberts 338, K; on fallen Picea branch with Hyphodonfia alufaria (Burt) Erikss., Bellever Forest, Dartmoor, South Devon (SX 6376), 20 Apr. 1991, P. Roberts 202 & 212, K; same location and substratum with Tylospora fibrillosa and hidiopsis oemifera (Obenv.) Woj.," 27 Apr. 1991, P. Roberts 213, K; same location and substratum, 6 May 1991, P. Roberts 225, K; same location and substratum, 1 Dec. 1991, P. Roberts 341, K; on fallen coniferous wood with Bofryobasidium subcoronaturn, Fernworthy Forest, Dartmoor, South Devon (SX 6583), 1 Mar. 1992, P. Roberts 366, K; on decaying wood, Yardley Chase, Northamptonshire (SP 8456), 6 Sep. 1991, P. Roberts 280, K; on Pinus log, New Forest, Wiltshire, Mar. 1991, A.Henrici, K; on fallen wood, Mickleham, Surrey (TQ 1753), 1 7 July 1991, A. Henrici, K.
Table 2. Tulasnella tomaculum: spore measurements (pm) from prints of the collections cited
Holotype P. Roberts 118 P. Roberts202 P. Roberts212 P. Roberts213 P. Roberts225 P. Roberts238 P. Roberts239 P. Roberts280 P. Roberts338 P. Roberts341 P. Roberts366 A. Henrici3/9l A. Henrici 7/91
(4.5-)5.06.5 x 2.5-3.0 4.5-70 x 2.5-3.0 (45-)5.0-65(-7.0) x 3.0-3.5 (4.0-)45-60(-5.5) x (2.5-)3.0(-3.5) (45-)5.0-65(-7.0) x 2.5-3.0 4.040 x 2.5-3.0 (4.0-)45+0(-65) x (2.0-)2.5(-3.0) 4.0-6.0 x 2.5-3.0 (4.5-)5.0-6.0(-5.5)x 2.5-3.0 40-7.5 x 2.0-2.5(-3.0) 40-60(-5.5) x 2.5-3.0 4.5-6.5(-7.0) x 2.5-3.0 (3.5-)5.0-75(-8.0) x (20-)2.5-3.0 4.540(-6.5) x 2.5-3.0
Tulasnella fornaculum is easily recognized b y its comparatively wide, unclamped hyphae and short, squat, weakly allantoid spores. Unlike T. allantospom, these spores normally have obtuse rather than tapered ends. Apart from its allantoid spores, T. fomaculum is identical to the ellipsoid-spored T . eichleriana Bres., and almost as common. So much so, that the collections cited above represent n o more than 5-10% of the total seen. It is surprising that such a common Tulasnella species should have remained undescribed until now, but it may well be that earlier collections have been referred t o T . allanfospora.
Tulasnella danica Hauersl., Friesia 11 (5): 275 (1979). (Fig. 4) ~ ~ ~ invisible i d to i the ~naked ~ eye ~ or~ ~nconsp~cuous, Fig. 4. Tulasnella danica. A, Mature 2-spored basidium (partly e h s e d , ceraceous, greyish, Hyphae unclamped or with rare, collapsed), hyphae bearing 4-spored basidia, and spores from ~ r i n t (P. Roberts 257); B, young 2 and 3-spored basidia, tangled basidiascattered 'lamps On h~phae (' Roberts 257); bearing hyphae, and spores from (P, Roberts 115), generally rather thin, ca 1.0-3.0 pm diam., but occasionally swollen to 4.0 pm or more, with distinct refractive walls in older specimens; forming a strongly branched, interlaced and entangled, basidia-bearing hymenium with hequent sterile UP t o 39.0 Pm o r more. Spores normally strongly allantoid and hyphal ends. Basidia normally sphaero-pedunculate, rather at each occas~ona~~y ellipsoid, ca 6.5-7.5 Pm wide, quickly collapsing (7.0-)90-15.0(-17-5) x (2.5-)3.0-4.0 vm when notionally in o n maturity. Sferigmata normally 2 per basidiurn, sometimes straightened out, 1-4, unusually widely spaced, for a 10% while remaining globose or ovoid, ca 4.0-6.5 pm wide, variously elongating
Specimens examined: on fallen Fraxinw branch near (but not in) Myxarium nucleatum Wallr., Orley Common, South Devon (SX
Peter Roberts 82661, 30 Dec. 1990, P. Roberts 115, K; same location, on fallen stick with Helicogloea lagerheimii and Xenasma pulverulenfum (Litsch.)Donk, 19 Jan. 1991, P. Roberts 130, K; same location, on fallen wood with Sisfotrema brinkrnannii (Bres.) Erikss., 5 Jan. 1991, P. Roberts 135, K; on decorticated fallen branch, probably Ulmus, Watcombe, Torquay, South Devon (SX 9267), 20 June 1991, P. Roberts 244, K; on fallen branch with Achroomyces vestifw (Bourd. & Galz.) Woj. and Tulasnella sp., Lincombe Slopes, Torquay, South Devon (SX 9363), 28 June 1991, P. Roberts 257, K.
Tulasnella danica is easily distinguished from other allantoidspored species by its unusual basidia and widely spaced sterigmata, its tangle-forming hymeniurn, hyphae lacking clamp-connexions (except rarely on the basal hyphae), and large, often deeply angled spores. As already noted by Hauerslev (1979), the basidia and sterigmata are identical to the Australian species, T. cruciata Warcup & Talbot, which has been isolated from orchids. A second Australian species, T. irregularis Warcup & Talbot, also belongs to this atypical group. All three species produce more or less sphaero-pedunculate basidia with widely spaced sterigmata, which at the extreme (in T . irregularis) can be equatorially distributed around the basidium. In Devon collections of T . danica the sterigmata, still remaining subglobose or ovoid, become enlarged whilst the basidium collapses, to the point where they are often difficult to distinguish from young basidia. By the time the sterigmata elongate and produce spores, the basidium has almost always collapsed completely. Devon collections of T . danica also show a unique hyrnenial arrangement, in which the basidia-bearing hyphal branches become increasingly interwoven and tangled, at the same time producing a substantial number of short, sterile hyphal ends. Fig. 4b shows the beginnings of such a tangle. The type collection of T . danica was found in the hymeniurn of Myxarium nucleatum, and on that basis has been keyed out as an intrahymenial parasite by Jiilich (1984). The Devon collections show that it is perfectly capable of forming independent fruit bodies. These collections represent the first records of T. danica in Britain.
Tulasnella saveloides P. Roberts, sp. nov.
(Figs 5, 6)
Etym.: from the English saveloy = a kind of sausage, with reference to the allantoid spores
Basidiomafa resupinata, inconspicua vel invisibilia licet observetur sine lente, ceraceo-gelatinosa, violaceo tincta. Hyphae non fibulatae, ca 2.5-3.5 Frn latae. Basidia fasciculata, clavata vel subglobosa, ca 5.5-7.0 um lata. Sterigmata 4, ellipsoidea vel fusoidea, saepe elongata. Sporae allantoideae, ca 65-8.5(-10.5) x 2.5-3.5 urn. Ad ligna putrida, Great Plantation, Bovey Tracey, South Devon, Anglia, 23 Mar. 1991, P. Roberts 142, holotypus, K.
Basidiomes often inconspicuous or invisible. When visible, effused, ceraceous or subgelatinous, violaceous. Hyphae unclamped, ca 2.5-3.5 ym wide, forming an open, sparsely branched, basidia-bearing hymenium in young specimens. In older specimens, basidia develop in clusters from single or contiguous sub-basidial cells. Conidia-like and gloeocystidialike cells seen in some specimens. Basidia varying hom subglobose to clavate, occasionally somewhat stipitate, ca
5.5-7.0 ym wide, forming in clusters (older specimens only) with old, collapsed, hyaline basidia conspicuously attached. Sterigmata 4 per basidium, ellipsoid to hsiform, ca 3.5-4.0 ym diam. at the widest point, variously elongating up to 26.5 ym or more. Spores strongly allantoid, often virtually rightangled, usually narrowing towards the apiculus, ca 6-5-8.5(-10.5) x 2.5-3.5 ym, replicating in mounts. 6.5-8.5 x 2-5-3.0 ym (P. Roberts 114); 7.5-9.0(-10.5) x 3-5(-4.0) ym (P. Roberts 131); 6.5-8.5 x 2.5-3-0 ym (P. Roberts 142, holotype); 6.5-8.0 x 2.5-3.0 ym (P. Roberts 313); 7.0-12.0 x 2.5-3.0(-3.5) ym (P. Roberts 329).
Specimens examined: on decorticated fallen C o y l w branch, Great Plantation, Bovey Tracey, South Devon (SX 8275), 27 Dec. 1990, P. Roberts 114, K; same location, on decorticated fallen Salix branch with clamped Tulasnella sp., 23 Mar. 1991, P. Roberts 142, holotype, K; on fallen branch, probably Fraxinus, with Achroomyces vestifus, Orley Common, South Devon (SX 8266), 26 Jan. 1991, P. Roberts 131, K ; same location, on fallen coniferous fence post with Tulasnella pennacra and Xenosperma ludibundum, 27 Oct. 1991, P. Roberts 313, K; same location, on decorticated Ulmus log, 23 Nov. 1991, P. Roberts 329, K. Tuhsnella saveloides can be recognized by its entirely unclamped hyphae, a hymenial growth pattern which produces basidia in clusters, and its comparatively wide, usually deeply angled spores similar to those of T . danica, but consistently shorter. The hymenial growth pattern of T. saveloides is not unique and has been found in a number of globose and ellipsoidspored species collected in Devon. When immature, these species generally produce very open growth with comparatively little hyphal branching. As a consequence, the terminal basidia are well-spaced and rather distant from each other. As the fruit body matures, new basidia are produced from the original sub-basidial cells (or new, contiguous subbasidial cells) gradually forming a cluster, with the old collapsed basidia remaining as hyaline shells. In old specimens, the hyphae up to and including the sub-basidial cells develop rather thickened, refractive walls. In squash mounts of these older specimens, the basidial clusters frequently become detached (see Fig. 5b). This cluster-forming hymenium appears to be distinct from the conventionally branching hymenium typical of the majority of Tulasnella species, though it is not an easy character to identify with certainty. In most Tulasnella species, new basidia-bearing branches proliferate from old ones, so that the hyrnenium continually expands, eventually becoming highly branched and compact. T.violea is a typical example. Tuhsnella saveloides appears to have two additional but inconstant features. Firstly, the type specimen is clearly producing coffin-shaped conidia or conidia-like cells as well as (and sometimes alongside) spore-bearing basidia (see Fig. 5a). This has not yet been observed in any other collection, or indeed in any other Tulasnefla species. Secondly, the collection P. Roberts 329 is ~roducinggloeocystidia-like cells which seem to be formed from two conjoined and often inflated basidia. Some have el lo wish, weakly refractive contents, others are hyaline (see Fig. 6). These are reminiscent of the much longer and highly irregular 'cystidia' found in the globose-spored T.cysfidiophora Hohn. & Litsch.
Allantoid-spored Tulasnella species from Devon
Fig. 5. Tulasnella saveloides. A, Young hyphal growth with coffin-shaped conidia-like cells, older cluster of basidia with collapsed basidia attached, and spores from print (holotype specimen); B, basidia showing variable sterigmata, detached cluster of mainly collapsed basidia (seen from above), replicating spores from mount, and spores from print (P. Roberts 114).
Tulasnella saveloides, as here defined, is a good and easily recognized species. But a number of anomalous collections have been seen with unclamped hyphae and weakly allantoid spores of similar size and shape to those of T . allantospora. These include Pearson's specimen (East Horsley, Surrey, 11 Apr. 1922, K) on which the original description of T. allantospora was based. This specimen shows signs of having a 'clustered' hymenium similar to T. saveloides, but other specimens from Devon are sparse and have a number of features which suggest they may not be conspecific. Following Julich (1984), all such unclamped collections might be assigned to T. rubropallens, though Bourdot & Galzin (1923) described this little-known species as having scattered clamp-connexions. Clearly further and better collections are required before any confident disposition can be made of these few remaining unclassified specimens.
Extra-limital allantoid-spored species
The following observations are based on published descriptions. Most of the species require reinvestigation and redescription, preferably based on hesh collections as well as herbarium material.
Tulasnella caroliniana (Olive) Olive, Mycologia 49: 665
(1957). As originally described and illustrated, this American species appears to be a synonym of T. allanfospora (qv.). However, Olive (1957) later assigned a collection from Tahiti to T. caroliniana which lacks clamp-connexions and is clearly a different species, probably identical to T. rubropallens in the sense of Julich (1984). It requires reinvestigation. Tulasnella curvispora Donk, Persoonia 4: 263 (1966).
Donk (1966) briefly described this Dutch species, without benefit of illustration, as having clamped hyphae and spores measuring 10-14 x 3-4 um. It is supposedly based on Rogers' (1933) interpretation of T . rutilans (Johan-Olsen ex Bref.) Juel. However, Rogers' wide and varied species concepts are such that American specimens should be re-examined and compared with European ones. T. curvispora should be recognizable by its long spores, but requires a full redescription. Tulasnella rogersii Olive, Mycologia 43: 688 (1951).
This American taxon is a nomen confmum composed of a clamped and unclamped species. As described and illustrated
Peter Roberts
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b y Olive (1951), the two species resemble T. allantospora and its unclamped counterpart. However T. rogersii is said to have very variable ellipsoid, clavate, or tubular gloeocystidia. These look similar t o the gloeocystidia-like cells described under T. saveloides above. Whether they have any taxonomic value is unclear, and both the species making up T. rogersii require reinvestigation.
T.rufilans is best regarded as a nomen dubium. Tulasnella rufilans as interpreted b y Rogers (1933) and subsequent American authors is said t o be T. curvispora (see note above). T. rufilans, as described and illustrated b y Christiansen (1959), appears to be T. danica. similar modern collections,
Tulasnella mbropallens Bourd. & Galz., Bull. Soc. Mycol. Fr. 39: 264 (1923). According t o Bourdot & Galzin (1923), this French species has scattered clamp-connexions and spores measuring 7.5-9 x 3-4 vm. Rogers (1933), followed b y most subsequent authors, placed it in synonymy with T. allantospora as suggested by Bourdot & Galzin (1928) themselves. However, Jiilich (1984) has resurrected 7. rubropallens as an unclamped species (see notes under T.allantospora and T. saveloides above). It clearly requires a full redescription.
Tulasnella rutilans (Johan-Olsen ex Bref.) Juel, Bih. Svensk. Vet.-Akad. Handl. 2 3 (3:12): 22 (1897). Brefeld (1889) illustrated this European species as having a conventionally branching hymenium with scattered clampconnexions, but the spore size can only be guessed at. Brefeld's measurements are '0.008 mm Lange und 0-016 mm Lange' (sic) which, even allowing for a misprint, is nonsense. If the illustrations are accurate, then (assuming the basidia are of average size, as suggested b y Brefeld) the weakly allantoid, and somewhat tapered spores should measure around n 12.0-13.0 x 3.0-3.5 Mm. This is somewhat longer than those 10 pm of T. rubro~allens~ a species also described as having scattered Fig. 6. Tulasnella saveloides. Clusters of basidia with gloeocystidiacells, and spores from print clamp-connexions, but comparable t o the clamped T. like cells, individual gloe~c~stidia-like curvispora. However, in the absence of a type specimen or any (P.Roberts 329).
Key to accepted allantoid-spored Tulasnella species 1 Clamp-cornexions present at all septa .
. . . . . 5 1 Clamp-cornexions absent, or scattered on basal hyphae only . . . 2 2 Spores mostly 90-15.0 um long, strongly angular. Basidia often two-spored. Sterigmata widely-spaced. Clamp-connexions rarely present on basal hyphae
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2 Spores mostly under 10.0 um long . . . . 3 Spores squat, mostly 40-70 um long, weakly allantoid. Hyphae always undamped 3 Spores mostly 7.0-90 um long, weakly allantoid or not .
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T. danica . 3 T. fornaculum 4
Spores weakly allantoid, tapering towards apiculus. Scattered clamp-connexions possible. Ill-defined species or species complex 7. rubropallens 4 Spores strongly angular. Basidia often in clusters. Hyphae undamped . . . . 7.saveloides 5 Spores mostly 10.0-140 um long . . . . . T. curvispora 5 Spores shorter, mostly under 10.0 um long . . . . . . . 6 6 Hyphae thin, sparse, 05-1.5 um wide. Basidia often unusually narrow and elongated. Fruitbody often (always?) no more than a . . . . . T. permacra few trailing, basidia-bearing hyphae 6 Hyphae 2.0-3.0 urn wide, normally forming an extensively branched and easily visible hymenium . . . T. allantospora 4
Thanks t o D r D. N. Pegler for facilities to examine specimens at the Royal Botanic Gardens, Kew.
REFERENCES Bourdot, H. & Galzin, A. (1923). Heterobasidieae nondum descriptae. Bulletin de la Sociik' mycologique de France 39, 261-266. Bourdot, H. & Galzin, A. (1928). Hyme'nomydtes de France. Sceaux.
Brefeld, 0. (1889). Die Gattung Pachysterigrna. Untersuchur~ggena m dem Gesamtgebiet der Mykologie 8 , 5-7. Christiansen, M. P. (1959). Danish resupinate fungi. 1. Ascomycetes and Heterobasidiornycetes. Dansk Botanisk Arkiv 19, 5-55. Donk, M. A. (1966). Cheddist of European hymenornycetous heterobasidiae. Persoonia 4, 145-335. Hauerslev, K. (1979). New or rare resupinate fungi horn Denmark. Friesia 11, 2 72-280.
Allantoid-spored Tulasnella species from Devon Jiilich, W. (1984). Die Nichtblatterpilze, Gallertpilze und Bauchpilze. In Kleine Kryptogamenfiora (ed. Gams, H.)I1 b / l . Stuttgart. Olive, L. S. (1951).Taxonomic notes on Louisiana fungi. 111. Additions to the Trernellales. Mycologia 43, 6 7 7 4 9 0 . Olive, L. S. (1953). New or noteworthy species of Tremellales from the Southern Appalachians. 11. Bulletin of the Torrey Botanical Club 80, 33-42.
Olive, L. S. (1957). Tulasnellaceae of Tahiti. A revision of the family. Mycologia 49, 6 6 3 4 7 9 . (Accepted 2 July 1992)
Roberts, P. (1992).Spiral-spored Tulasnella species from Devon and the New Forest. Mycological Research 96, 233-236. Rogers, D. P. (1933). A taxonomic review of the Tulasnellaceae. Annales Mycologici 31, 181-203.
Wakefield, E. M. & Pearson, A. A. ( 1923). Some additional records of Surrey resupinate hyrnenornycetes. Transactions of the British Mycological Society 8, 216-221.
Mycol. Res. 97 (2): 213-220 (1993) Printed in Great Britain
Allantoid-spored Tulasnella species from Devon
PETER ROBERTS 36 Western Road, Torquay, Devon TQI 4RL
Tulasnella permacra, T. saveloides, and T. fomaculum are described as new. T. danica is described and recorded as new to Britain. T. allantospora is redescribed with reference to the holotype. Extra-limital species are briefly noted and a key is provided to accepted taxa.
Extensive collecting 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. More emphasis is placed on hyphal, spore, and hymenial morphology and less on fruit-body size, colour, texture, and habitat. An initial paper (Roberts, 1992) looked at four species with spiral spores. This second contribution examines five allantoidspored species found in Devon, with reference to collections made elsewhere in Britain plus published descriptions of overseas species. Tulasnella appears to be a genus rich in species and it is highly unlikely that a few woodlands in South Devon will have yielded up more than a small proportion of them. Care should therefore be taken before automatically assigning collections from elsewhere to the following species.
Tulasnella allantospora Wakef. & Pears., Trans. Br. Mycol. SOC.8 : 220 (1923) (Fig. I) Gloeotulasnella caroliniana Olive, Bull. Torrey Bof. Club 80 (1): 41 (1953).
Basidiomes often invisible to the naked eye, or forming no more than a faint, dull bloom. When visible, effused, ceraceous or subgelatinous, violaceous grey. Hyphae clamped at all septa, ca 2.0-3.0 pm diam., forming an extensively branched, basidia-bearing hymenium. Basidia variously clavate to subglobose, ca 4-0-6.5 vm wide, with or without a short or long stalk, clamped at the base. Sterigmata 4 per basidium, ellipsoid becoming fusiform, ca 3.0-4.0 pm diam. at widest point, variously extending up to 12.5 ym or more. Spores weakly allantoid, tapered at both ends, averaging 6.5-9.0(-10.0) x 2.5-3.0 pm (see Table I), replicating in mounts. Specimens examined: on fallen Pinus branch, Great Plantation, Bovey Tracey, South Devon (SX 8275), 12 Jan. 1991, P. Roberts 124, K; same location and substratum, 16 June 1991, P. Roberts 240, K; on decorticated Picea log with Tylospora jbrillosa (Burt) Donk and Tulasnella sp., Bellever Forest, Dartmoor, South Devon (SX 6376). 27
Apr. 1991, P. Roberts 220, K; same location, on fallen Picea branch, I Dec. 1991, P. Roberts 340, K; on fallen coniferous fence-post, Orley Common, South Devon (SX 8266), 9 Nov. 1991, P. Roberts 318, K; same location, on fallen deciduous branch, 30 Nov. 1991, P.Roberts 339, K ; on fallen branch, probably Picea, with Botyobasidium subcoronatum (Hohn. & Litsch.) Donk, Fernworthy Forest, Dartmoor, South Devon (SX 6583), 1 Mar. 1992, P. Roberts 365, K; on decorticated coniferous wood, East Horsley, Surrey, 4 Apr. 1922, A. Pearson, holotype, K.
Tulasnella alhnfospora has been a much-confused species ever since it was first described. Wakefield & Pearson (1923) based their type description on two specimens collected from the same locality a week apart. Unfortunately these represent two different species. As already pointed out by Rogers (1933), the earliest specimen (taken to be the holotype) possesses abundant and clearly visible clamp-connexions. The second collection, on which Wakefield & Pearson's description and illustration seem to be based, has similar spores but lacks clamp-connexions. As a result, T. allantospora has been variously interpreted in the literature and all records are suspect unless accompanied by a full description or voucher material. Based on the holotype, T. allantospora is a consistent and easily recognized species, distinguished by its conventionally branching hymenium with clamps at all septa, together with its weakly allantoid spores, the majority of which (in every print) taper towards each end, more particularly towards the apiculus. It has also proved to be one of the commonest Tulasnella species in Devon, and the collections cited above are just a representative sample of the total seen. T. interrogans P. Roberts is a closely related but less common species with spiral spores. Bourdot & Galzin (1923) originally described Tulasnella rubropallens Bourd. & Galz. as having similar spores (measuring 7.5-9 x 3-4 pm) but with only scattered clamp-connexions on the hyphae and none at all on the basidia. The hyphae and basidia were described as wider. Later (Bourdot & Galzin,
Allantoid-spored Tulasnella species from Devon
P-, A
Fig. 1. Tulasnefla allantospora. A, Basidium and spores from mount (holotype specimen); B, (compact fruit body) branching hyphae, basidia, and spores from print (P. Roberts 124); C, (more open fruit body) basidia, spores from print, and hyphae showing continued growth through the basidia (P. Roberts 318).
Table I . Tulasnella allantospora: spore measurements (pm) from prints of the collections cited plus relevant publications Holotype* P. Roberts 124 P. Roberts 220 P. Roberts 240 P. Roberts 318 P. Roberts 339 P. Roberts 340 P. Roberts 365 Jiilich (1984) Olive (1953)t * Measurements taken from a few scattered spores in a mount. The original published spore measurements almost certainly refer to a different, unclamped species t Published measurements for holotype of Gloeotulasnella caroliniana.
1928), they suggested their species might be conspecific with T. allantospora, a view shared by most subsequent authors. Jiilich (1984), however, has repositioned T. rubropallens as an entirely unclamped species with spores measuring 7-9(-10) x 2.4-3.4 pm, though on what basis is unclear. It urgently requires a modem redescription. Olive (1953) described an American species, Gloeofulasnella caroliniana Olive, which appears to have identical spores and is distinguishable from T. allanfospora only by its subgelatinous
fruit body. Since fruit-body texture has more to do with humidity than taxonomy, G. caroliniana is here proposed as a synonym of T. allantospora. Tulasnella permacra P. Roberts, sp. nov.
(Fig. 2)
Etym.: from the Latin, per plus macer = very meagre
Basidiomata resupinata, inconspicua vel invisibilia licet observetur sine lente. Hyphae fibulatae, angustae, 0.5-1.5 pm latae. Basidia stipitata, obovata vel clavata, angusta, ca 2.5-4.0 ym lata. Sferigmafa 4, fusoidea. Sporae allantoideae, T. allantospora proximae sed minores. ca 5.5-8.0 x 2.0-2'5 pm. Ad ligna ~utrida,Orley Common, Devon, Anglia, 19 Jan. 1991, P. Roberts 125, holotypus, K. Basidiomes resupinate, inconspicuous or invisible to the unaided eye. Hyphae clamped, unusually and consistently thin, ca 0.5-1.5 pm diam., sparsely branched, not seen to form a hymenium but trailing through and over the substratum, producing basidia individually on short side branches. Basidia narrowly clavate or ovoid, ca 2.5-4.0 pm wide, with a distinct stalk, clamped at the base. Basidia frequently appear elongated or narrowly fusiform as a result of apical hyphal outgrowths. Sterigmata 4 per basidium, ellipsoid, becoming widely fusiform, ca 3.0-4.5 pm diam. at the widest point, variously extending up to 8.5 pm. Spores similar to T. albntospora but generally shorter, narrower, and more strongly curved, ca
Peter Roberts
Fig. 2. Tulasnella permacra. A, Typically thin hyphae with ovoid and elongated basidia, plus spores from print (holotype specimen); B, basidia and spores from print (P. Roberts 252).
5.5-8.0 X 2.0-2.5 ym; replicating in mounts. (6-0-)6.58-0(-8.5) x (2-0-)2.5(-3-0) ym (P. Roberts 125, holotype); 5.5-8.0 x 2.0(-2.5) ym (P. Roberts 252); 5.0-6.0 x 2.02.5 ym (P. Roberts 312).
Specimens examined: on fallen decorticated Picea branch with Tulasnella violea (QuCl.) Bourd. & Galz., Stover Park, South Devon (SX 8375), 11 Nov. 1989, P. Roberts 101, K; on fallen Fraxinus branch, Orley Common, South Devon (SX 8266), 19 Jan. 1991, P. Roberts 125, holotype, K; same location, on fallen deciduous wood with Tulasnella sp. and Helicogloea lagerheimii Pat., 22 June 1991, P. Roberts 252, K; same location, on fallen coniferous fence post with Xenosperma ludibundtrm (Rogers & Liberta) Oberw. ex Julich and Iulasnella sp., 27 Oct. 1991, P. Roberts 312, K. Tulasnella pemacra is a cryptic but distinctive species, easily separated from T. allantospora b y its exceptionally thin, clamped hyphae, tiny and unusually narrow basidia, and rather smaller, narrower, and more strongly allantoid spores. The few collections made, all of which are cited above, consist solely of trailing, sparsely branched hyphae producing basidia singly o n short side branches. However, 7.permacra does not appear t o be an obligate hymenial parasite and there is n o intrinsic reason why it should not form a more compact hymenium when growth conditions permit. T h e exceptionally narrow basidia are a constant feature of the species. Their narrowness is exacerbated b y a tendency for hyphal growth t o continue through the basidium, emerging a t
Fig. 3. Tulasnella tomaculum. A, Basidia-bearing hyphae, replicating spore, basidia showing variable sterigmata, and spores from print (holotype specimen); B, young, open growth in the hymenium and spores from print (P. Roberts 341).
the apex (or occasionally laterally) as a papillate or snout-like outgrowth. Such basidial outgrowths are common in Tulasnelb species, and are particularly prevalent in young o r sparse collections, less evident or absent when a stable hymenium is formed. The formation of sterigmata usually prevents further hyphal growth from the basidium; where growth does continue, the basidium is normally left as a sterile cell or reduced t o little more than a swelling. Very occasionally immature sterigmata can be found o n such swellings (see T . allantospora, Fig. Ic), but none has yet been seen t o produce spores.
Tulasnella tomaculum P. Roberts, sp. nov.
(Fig. 3)
Etym.: from the Latin, fornaculum = a kind of small sausage, with reference t o the allantoid spores
Basidiomafa resupinata, inconspicua, ceraceo-gelatinosa, lilaceo-grisea tincta. Hyphae non fibulatae, ca 2.5-3.5 pm latae. Basidia clavata vel obovata, ca 4.0-5.5 pm lata. Sterigmata 4, fusoidea vel clavata. Sporae breviter allantoideae, ca 4.0-7.0 x 2.5-3.0 pm. Ad ligna putrida, Orley Common, South Devon, Anglia, 27 Dec. 1990, P. Roberts 119, holotypus, K. Basidiomes often inconspicuous o r invisible t o the naked eye. When visible, effused, resupinate, ceraceous t o subgelatinous,
216
Allantoid-spored Tulasnella species from Devon violaceous-grey. Hyphae unclamped, forming a conventionally branching and often compact basidia-bearing hymenium. Basal hyphae normally long and straight, somewhat thickwalled in mature specimens; subh~menial hyphae rather short-celled and slightly swollen, ca 2-5-3.5 pm diam. Basidia normally clavate, but frequently ovoid or subglobose, ca 4.0-5.5 pm wide, often with a poorly defined long or short stalk. Sferigmata 4 per basidium, variable but normally fusiform, sometimes elongated or clavate, ca 2.5-3.5 pm diam. at the widest point, variously extending- up. to 21.0 pm or more. Spores weakly allantoid, normally obtuse at each end, ca 4.0-7.0 x 2.5-3.0 pm (see Table 2), replicating in mounts. Specimens examined: on bark of Picea log, Great Plantation, Bovey Tracey, South Devon (SX 8275), 1Apr. 1989, P. Roberts 96, K; same location and substratum, 16 June 1991, P. Roberts 238 & 239, K; on fallen Picea branch, Stover Park, South Devon (SX 8375), 1Dec. 1990, P. Roberts 118, K; on decorticated deciduous branch, possibly C o y l w , Orley Common, South Devon (SX 8266), 27 Dec. 1990, P. Roberts 119, holotype, K; same location, on fallen deciduous wood, 30 Nov. 1991, P. Roberts 338, K; on fallen Picea branch with Hyphodonfia alufaria (Burt) Erikss., Bellever Forest, Dartmoor, South Devon (SX 6376), 20 Apr. 1991, P. Roberts 202 & 212, K; same location and substratum with Tylospora fibrillosa and hidiopsis oemifera (Obenv.) Woj.," 27 Apr. 1991, P. Roberts 213, K; same location and substratum, 6 May 1991, P. Roberts 225, K; same location and substratum, 1 Dec. 1991, P. Roberts 341, K; on fallen coniferous wood with Bofryobasidium subcoronaturn, Fernworthy Forest, Dartmoor, South Devon (SX 6583), 1 Mar. 1992, P. Roberts 366, K; on decaying wood, Yardley Chase, Northamptonshire (SP 8456), 6 Sep. 1991, P. Roberts 280, K; on Pinus log, New Forest, Wiltshire, Mar. 1991, A.Henrici, K; on fallen wood, Mickleham, Surrey (TQ 1753), 1 7 July 1991, A. Henrici, K.
Table 2. Tulasnella tomaculum: spore measurements (pm) from prints of the collections cited
Holotype P. Roberts 118 P. Roberts202 P. Roberts212 P. Roberts213 P. Roberts225 P. Roberts238 P. Roberts239 P. Roberts280 P. Roberts338 P. Roberts341 P. Roberts366 A. Henrici3/9l A. Henrici 7/91
(4.5-)5.06.5 x 2.5-3.0 4.5-70 x 2.5-3.0 (45-)5.0-65(-7.0) x 3.0-3.5 (4.0-)45-60(-5.5) x (2.5-)3.0(-3.5) (45-)5.0-65(-7.0) x 2.5-3.0 4.040 x 2.5-3.0 (4.0-)45+0(-65) x (2.0-)2.5(-3.0) 4.0-6.0 x 2.5-3.0 (4.5-)5.0-6.0(-5.5)x 2.5-3.0 40-7.5 x 2.0-2.5(-3.0) 40-60(-5.5) x 2.5-3.0 4.5-6.5(-7.0) x 2.5-3.0 (3.5-)5.0-75(-8.0) x (20-)2.5-3.0 4.540(-6.5) x 2.5-3.0
Tulasnella fornaculum is easily recognized b y its comparatively wide, unclamped hyphae and short, squat, weakly allantoid spores. Unlike T. allantospom, these spores normally have obtuse rather than tapered ends. Apart from its allantoid spores, T. fomaculum is identical to the ellipsoid-spored T . eichleriana Bres., and almost as common. So much so, that the collections cited above represent n o more than 5-10% of the total seen. It is surprising that such a common Tulasnella species should have remained undescribed until now, but it may well be that earlier collections have been referred t o T . allanfospora.
Tulasnella danica Hauersl., Friesia 11 (5): 275 (1979). (Fig. 4) ~ ~ ~ invisible i d to i the ~naked ~ eye ~ or~ ~nconsp~cuous, Fig. 4. Tulasnella danica. A, Mature 2-spored basidium (partly e h s e d , ceraceous, greyish, Hyphae unclamped or with rare, collapsed), hyphae bearing 4-spored basidia, and spores from ~ r i n t (P. Roberts 257); B, young 2 and 3-spored basidia, tangled basidiascattered 'lamps On h~phae (' Roberts 257); bearing hyphae, and spores from (P, Roberts 115), generally rather thin, ca 1.0-3.0 pm diam., but occasionally swollen to 4.0 pm or more, with distinct refractive walls in older specimens; forming a strongly branched, interlaced and entangled, basidia-bearing hymenium with hequent sterile UP t o 39.0 Pm o r more. Spores normally strongly allantoid and hyphal ends. Basidia normally sphaero-pedunculate, rather at each occas~ona~~y ellipsoid, ca 6.5-7.5 Pm wide, quickly collapsing (7.0-)90-15.0(-17-5) x (2.5-)3.0-4.0 vm when notionally in o n maturity. Sferigmata normally 2 per basidiurn, sometimes straightened out, 1-4, unusually widely spaced, for a 10% while remaining globose or ovoid, ca 4.0-6.5 pm wide, variously elongating
Specimens examined: on fallen Fraxinw branch near (but not in) Myxarium nucleatum Wallr., Orley Common, South Devon (SX
Peter Roberts 82661, 30 Dec. 1990, P. Roberts 115, K; same location, on fallen stick with Helicogloea lagerheimii and Xenasma pulverulenfum (Litsch.)Donk, 19 Jan. 1991, P. Roberts 130, K; same location, on fallen wood with Sisfotrema brinkrnannii (Bres.) Erikss., 5 Jan. 1991, P. Roberts 135, K; on decorticated fallen branch, probably Ulmus, Watcombe, Torquay, South Devon (SX 9267), 20 June 1991, P. Roberts 244, K; on fallen branch with Achroomyces vestifw (Bourd. & Galz.) Woj. and Tulasnella sp., Lincombe Slopes, Torquay, South Devon (SX 9363), 28 June 1991, P. Roberts 257, K.
Tulasnella danica is easily distinguished from other allantoidspored species by its unusual basidia and widely spaced sterigmata, its tangle-forming hymeniurn, hyphae lacking clamp-connexions (except rarely on the basal hyphae), and large, often deeply angled spores. As already noted by Hauerslev (1979), the basidia and sterigmata are identical to the Australian species, T. cruciata Warcup & Talbot, which has been isolated from orchids. A second Australian species, T. irregularis Warcup & Talbot, also belongs to this atypical group. All three species produce more or less sphaero-pedunculate basidia with widely spaced sterigmata, which at the extreme (in T . irregularis) can be equatorially distributed around the basidium. In Devon collections of T . danica the sterigmata, still remaining subglobose or ovoid, become enlarged whilst the basidium collapses, to the point where they are often difficult to distinguish from young basidia. By the time the sterigmata elongate and produce spores, the basidium has almost always collapsed completely. Devon collections of T . danica also show a unique hyrnenial arrangement, in which the basidia-bearing hyphal branches become increasingly interwoven and tangled, at the same time producing a substantial number of short, sterile hyphal ends. Fig. 4b shows the beginnings of such a tangle. The type collection of T . danica was found in the hymeniurn of Myxarium nucleatum, and on that basis has been keyed out as an intrahymenial parasite by Jiilich (1984). The Devon collections show that it is perfectly capable of forming independent fruit bodies. These collections represent the first records of T. danica in Britain.
Tulasnella saveloides P. Roberts, sp. nov.
(Figs 5, 6)
Etym.: from the English saveloy = a kind of sausage, with reference to the allantoid spores
Basidiomafa resupinata, inconspicua vel invisibilia licet observetur sine lente, ceraceo-gelatinosa, violaceo tincta. Hyphae non fibulatae, ca 2.5-3.5 Frn latae. Basidia fasciculata, clavata vel subglobosa, ca 5.5-7.0 um lata. Sterigmata 4, ellipsoidea vel fusoidea, saepe elongata. Sporae allantoideae, ca 65-8.5(-10.5) x 2.5-3.5 urn. Ad ligna putrida, Great Plantation, Bovey Tracey, South Devon, Anglia, 23 Mar. 1991, P. Roberts 142, holotypus, K.
Basidiomes often inconspicuous or invisible. When visible, effused, ceraceous or subgelatinous, violaceous. Hyphae unclamped, ca 2.5-3.5 ym wide, forming an open, sparsely branched, basidia-bearing hymenium in young specimens. In older specimens, basidia develop in clusters from single or contiguous sub-basidial cells. Conidia-like and gloeocystidialike cells seen in some specimens. Basidia varying hom subglobose to clavate, occasionally somewhat stipitate, ca
5.5-7.0 ym wide, forming in clusters (older specimens only) with old, collapsed, hyaline basidia conspicuously attached. Sterigmata 4 per basidium, ellipsoid to hsiform, ca 3.5-4.0 ym diam. at the widest point, variously elongating up to 26.5 ym or more. Spores strongly allantoid, often virtually rightangled, usually narrowing towards the apiculus, ca 6-5-8.5(-10.5) x 2.5-3.5 ym, replicating in mounts. 6.5-8.5 x 2-5-3.0 ym (P. Roberts 114); 7.5-9.0(-10.5) x 3-5(-4.0) ym (P. Roberts 131); 6.5-8.5 x 2.5-3-0 ym (P. Roberts 142, holotype); 6.5-8.0 x 2.5-3.0 ym (P. Roberts 313); 7.0-12.0 x 2.5-3.0(-3.5) ym (P. Roberts 329).
Specimens examined: on decorticated fallen C o y l w branch, Great Plantation, Bovey Tracey, South Devon (SX 8275), 27 Dec. 1990, P. Roberts 114, K; same location, on decorticated fallen Salix branch with clamped Tulasnella sp., 23 Mar. 1991, P. Roberts 142, holotype, K; on fallen branch, probably Fraxinus, with Achroomyces vestifus, Orley Common, South Devon (SX 8266), 26 Jan. 1991, P. Roberts 131, K ; same location, on fallen coniferous fence post with Tulasnella pennacra and Xenosperma ludibundum, 27 Oct. 1991, P. Roberts 313, K; same location, on decorticated Ulmus log, 23 Nov. 1991, P. Roberts 329, K. Tuhsnella saveloides can be recognized by its entirely unclamped hyphae, a hymenial growth pattern which produces basidia in clusters, and its comparatively wide, usually deeply angled spores similar to those of T . danica, but consistently shorter. The hymenial growth pattern of T. saveloides is not unique and has been found in a number of globose and ellipsoidspored species collected in Devon. When immature, these species generally produce very open growth with comparatively little hyphal branching. As a consequence, the terminal basidia are well-spaced and rather distant from each other. As the fruit body matures, new basidia are produced from the original sub-basidial cells (or new, contiguous subbasidial cells) gradually forming a cluster, with the old collapsed basidia remaining as hyaline shells. In old specimens, the hyphae up to and including the sub-basidial cells develop rather thickened, refractive walls. In squash mounts of these older specimens, the basidial clusters frequently become detached (see Fig. 5b). This cluster-forming hymenium appears to be distinct from the conventionally branching hymenium typical of the majority of Tulasnella species, though it is not an easy character to identify with certainty. In most Tulasnella species, new basidia-bearing branches proliferate from old ones, so that the hyrnenium continually expands, eventually becoming highly branched and compact. T.violea is a typical example. Tuhsnella saveloides appears to have two additional but inconstant features. Firstly, the type specimen is clearly producing coffin-shaped conidia or conidia-like cells as well as (and sometimes alongside) spore-bearing basidia (see Fig. 5a). This has not yet been observed in any other collection, or indeed in any other Tulasnefla species. Secondly, the collection P. Roberts 329 is ~roducinggloeocystidia-like cells which seem to be formed from two conjoined and often inflated basidia. Some have el lo wish, weakly refractive contents, others are hyaline (see Fig. 6). These are reminiscent of the much longer and highly irregular 'cystidia' found in the globose-spored T.cysfidiophora Hohn. & Litsch.
Allantoid-spored Tulasnella species from Devon
Fig. 5. Tulasnella saveloides. A, Young hyphal growth with coffin-shaped conidia-like cells, older cluster of basidia with collapsed basidia attached, and spores from print (holotype specimen); B, basidia showing variable sterigmata, detached cluster of mainly collapsed basidia (seen from above), replicating spores from mount, and spores from print (P. Roberts 114).
Tulasnella saveloides, as here defined, is a good and easily recognized species. But a number of anomalous collections have been seen with unclamped hyphae and weakly allantoid spores of similar size and shape to those of T . allantospora. These include Pearson's specimen (East Horsley, Surrey, 11 Apr. 1922, K) on which the original description of T. allantospora was based. This specimen shows signs of having a 'clustered' hymenium similar to T. saveloides, but other specimens from Devon are sparse and have a number of features which suggest they may not be conspecific. Following Julich (1984), all such unclamped collections might be assigned to T. rubropallens, though Bourdot & Galzin (1923) described this little-known species as having scattered clamp-connexions. Clearly further and better collections are required before any confident disposition can be made of these few remaining unclassified specimens.
Extra-limital allantoid-spored species
The following observations are based on published descriptions. Most of the species require reinvestigation and redescription, preferably based on hesh collections as well as herbarium material.
Tulasnella caroliniana (Olive) Olive, Mycologia 49: 665
(1957). As originally described and illustrated, this American species appears to be a synonym of T. allanfospora (qv.). However, Olive (1957) later assigned a collection from Tahiti to T. caroliniana which lacks clamp-connexions and is clearly a different species, probably identical to T. rubropallens in the sense of Julich (1984). It requires reinvestigation. Tulasnella curvispora Donk, Persoonia 4: 263 (1966).
Donk (1966) briefly described this Dutch species, without benefit of illustration, as having clamped hyphae and spores measuring 10-14 x 3-4 um. It is supposedly based on Rogers' (1933) interpretation of T . rutilans (Johan-Olsen ex Bref.) Juel. However, Rogers' wide and varied species concepts are such that American specimens should be re-examined and compared with European ones. T. curvispora should be recognizable by its long spores, but requires a full redescription. Tulasnella rogersii Olive, Mycologia 43: 688 (1951).
This American taxon is a nomen confmum composed of a clamped and unclamped species. As described and illustrated
Peter Roberts
219
b y Olive (1951), the two species resemble T. allantospora and its unclamped counterpart. However T. rogersii is said to have very variable ellipsoid, clavate, or tubular gloeocystidia. These look similar t o the gloeocystidia-like cells described under T. saveloides above. Whether they have any taxonomic value is unclear, and both the species making up T. rogersii require reinvestigation.
T.rufilans is best regarded as a nomen dubium. Tulasnella rufilans as interpreted b y Rogers (1933) and subsequent American authors is said t o be T. curvispora (see note above). T. rufilans, as described and illustrated b y Christiansen (1959), appears to be T. danica. similar modern collections,
Tulasnella mbropallens Bourd. & Galz., Bull. Soc. Mycol. Fr. 39: 264 (1923). According t o Bourdot & Galzin (1923), this French species has scattered clamp-connexions and spores measuring 7.5-9 x 3-4 vm. Rogers (1933), followed b y most subsequent authors, placed it in synonymy with T. allantospora as suggested by Bourdot & Galzin (1928) themselves. However, Jiilich (1984) has resurrected 7. rubropallens as an unclamped species (see notes under T.allantospora and T. saveloides above). It clearly requires a full redescription.
Tulasnella rutilans (Johan-Olsen ex Bref.) Juel, Bih. Svensk. Vet.-Akad. Handl. 2 3 (3:12): 22 (1897). Brefeld (1889) illustrated this European species as having a conventionally branching hymenium with scattered clampconnexions, but the spore size can only be guessed at. Brefeld's measurements are '0.008 mm Lange und 0-016 mm Lange' (sic) which, even allowing for a misprint, is nonsense. If the illustrations are accurate, then (assuming the basidia are of average size, as suggested b y Brefeld) the weakly allantoid, and somewhat tapered spores should measure around n 12.0-13.0 x 3.0-3.5 Mm. This is somewhat longer than those 10 pm of T. rubro~allens~ a species also described as having scattered Fig. 6. Tulasnella saveloides. Clusters of basidia with gloeocystidiacells, and spores from print clamp-connexions, but comparable t o the clamped T. like cells, individual gloe~c~stidia-like curvispora. However, in the absence of a type specimen or any (P.Roberts 329).
Key to accepted allantoid-spored Tulasnella species 1 Clamp-cornexions present at all septa .
. . . . . 5 1 Clamp-cornexions absent, or scattered on basal hyphae only . . . 2 2 Spores mostly 90-15.0 um long, strongly angular. Basidia often two-spored. Sterigmata widely-spaced. Clamp-connexions rarely present on basal hyphae
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2 Spores mostly under 10.0 um long . . . . 3 Spores squat, mostly 40-70 um long, weakly allantoid. Hyphae always undamped 3 Spores mostly 7.0-90 um long, weakly allantoid or not .
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T. danica . 3 T. fornaculum 4
Spores weakly allantoid, tapering towards apiculus. Scattered clamp-connexions possible. Ill-defined species or species complex 7. rubropallens 4 Spores strongly angular. Basidia often in clusters. Hyphae undamped . . . . 7.saveloides 5 Spores mostly 10.0-140 um long . . . . . T. curvispora 5 Spores shorter, mostly under 10.0 um long . . . . . . . 6 6 Hyphae thin, sparse, 05-1.5 um wide. Basidia often unusually narrow and elongated. Fruitbody often (always?) no more than a . . . . . T. permacra few trailing, basidia-bearing hyphae 6 Hyphae 2.0-3.0 urn wide, normally forming an extensively branched and easily visible hymenium . . . T. allantospora 4
Thanks t o D r D. N. Pegler for facilities to examine specimens at the Royal Botanic Gardens, Kew.
REFERENCES Bourdot, H. & Galzin, A. (1923). Heterobasidieae nondum descriptae. Bulletin de la Sociik' mycologique de France 39, 261-266. Bourdot, H. & Galzin, A. (1928). Hyme'nomydtes de France. Sceaux.
Brefeld, 0. (1889). Die Gattung Pachysterigrna. Untersuchur~ggena m dem Gesamtgebiet der Mykologie 8 , 5-7. Christiansen, M. P. (1959). Danish resupinate fungi. 1. Ascomycetes and Heterobasidiornycetes. Dansk Botanisk Arkiv 19, 5-55. Donk, M. A. (1966). Cheddist of European hymenornycetous heterobasidiae. Persoonia 4, 145-335. Hauerslev, K. (1979). New or rare resupinate fungi horn Denmark. Friesia 11, 2 72-280.
Allantoid-spored Tulasnella species from Devon Jiilich, W. (1984). Die Nichtblatterpilze, Gallertpilze und Bauchpilze. In Kleine Kryptogamenfiora (ed. Gams, H.)I1 b / l . Stuttgart. Olive, L. S. (1951).Taxonomic notes on Louisiana fungi. 111. Additions to the Trernellales. Mycologia 43, 6 7 7 4 9 0 . Olive, L. S. (1953). New or noteworthy species of Tremellales from the Southern Appalachians. 11. Bulletin of the Torrey Botanical Club 80, 33-42.
Olive, L. S. (1957). Tulasnellaceae of Tahiti. A revision of the family. Mycologia 49, 6 6 3 4 7 9 . (Accepted 2 July 1992)
Roberts, P. (1992).Spiral-spored Tulasnella species from Devon and the New Forest. Mycological Research 96, 233-236. Rogers, D. P. (1933). A taxonomic review of the Tulasnellaceae. Annales Mycologici 31, 181-203.
Wakefield, E. M. & Pearson, A. A. ( 1923). Some additional records of Surrey resupinate hyrnenornycetes. Transactions of the British Mycological Society 8, 216-221.