• Br. mycol. Soc. 74 (2) 297-307 (1980) Printed in Great Britain

THE MORENOINA IN BRITAIN

BY J. PAMELA ELLIS 31 Marlborough Road, Southwold, Suffolk

The type species of Morenoina Theiss. is illustrated; nine new British species and two from Europe, not yet found in Britain, are described. Aulographum hederae is figured for com­ parison.

Morenoina Theiss. is in the family Asterinaceae, Ascospores 1-septate, at first hyaline and guttulate, order Hemisphaeriales of the Loculoascomycetes. but at maturity often becoming faintly brown, lose The genus has the following characters: their guttules and their walls may become minutely Colonies grow on decaying plant debris and are rough; in all cases their ends are rounded and the composed of elongated, black, often branched upper cell is a little wider than the lower one. ascomata usually less than 1 mm in length and must Conidial states have been observed in three species. be sought with a hand-lens. Mycelium superficial, Conidia produced in circular, radiately scutellate usually fine, and sometimes very sparse and easily pycnothyria which: have been seen to be connected rubbed off in older colonies, but may be more directly by hyphae to the thyriothecia. robust and quite abundant in a few species. Fig. 1 illustrates Morenoina antarctica (Speg.) Thyriothecia start as small circular radiate shields Theiss., the type species of the genus. This figure and develop in two opposite directions to become is taken from a collection in Herb.K examined and elongated; sometimes they branch to become confirmed by Prof. E. Muller and compared by him Y-shaped, X-shaped or irregularly lobed. There is with the type collection. Muller & von Arx (1962) no hypostroma. Scutellum brown, darker towards described this species fully. A genus sometimes the centre and composed of radiately-arranged confused with Morenoina is Aulographum Lib. This more-or-Iess rectangular cells. The cells at the is characterized by having an elongated scutellum margin vary considerably in different species and composed of inordinately arranged cells. Muller & this character is best seen by reference to the von Arx (1962) showed that the type species, illustrations. The scutellum splits longitudinally at A. hederae Lib. (Fig. 2), has a hypostroma con­ maturity to expose the abundant asci. Asci sisting of an extensive plate of subcuticular hyphae arranged vertically beneath the scutellum with their beneath the thyriothecium. Luttrell (1973) puts bases attached to a very thin basal plate which is Aulographum into a separate family of the Hemi­ composed of hyaline or pale brown radially sphaeriales, the Aulographaceae. The photographs arranged cells mirroring the upper wall. The illustrating the structure of the thyriothecium of bitunicate asci are variable in shape, spherical, ovoid A. hederae are taken from part of the type collection or clavate, this often depending on their stage of in Herb.K .This species name was validly published development, and contain eight variously arranged by Libert (1834). A later name for this is ascospores. The asci are embedded in a slimy mass A. vagum Desmazieres (1843). and no pseudoparaphyses have been observed.

KEY TO THE SPECIES OF MORENOINA Ascospores up to 18 fim long 1 Ascospores not more than 15 fim long ...... 2 1. Ascospores smooth, on Rubus ...... clarkii Ascospores closely echinulate, on Arundinaria ...... arundinariae 2. Ascospores 9-15 fim long ...... 3 Ascospores never more than 10 fim long ...... 5 3- Conidia 5-7 x 1-5-2 fim present in pycnothyria, on Carex, Cladium and Juncus . . . • paludosa Conidia absent 4 4. Branches of thyriothecia 150-200 fim wide, ascospores 10-14 x 4-5 fim, on Arundinaria • • websteri Branches of thyriothecia 80-120 fim wide, ascospores 12-15 x 4-5-5 fim, on Carex and Juncus . fimbriata 5. Ascospores not more than 7 fim long ...... epilobii* Ascospores more than 7 fim and less than to fim long ...... g 6. Conidia present in pycnothyria J Conidia absent ...... g 7. Ascospores 8-iO x 3-4 fim, conidia 3-4 x 0-5-1 fim, on Phalaris and Phragmites • • phragmitidis Ascospores 8-10 x 2-5-3-5 fim, conidia 2-3 x 0-5-1 fim, on Rhododendron and Vaccinium . rhododendri 8. Thyriothecia not more than 250 fim long, on Carex, Cladium and Juncus .... minuta Thyriothecia mostly more than 250 fim long

* European species not yet found in Britain.

Fig. 1. Morenoina antarctica. (A) Thyriothecia ( x 100); (B) edge of scutellum; (C) ascospores.

Fig. 2. Aulographum hederae. (A) Thyriothecium ( x 250); (B) part of thyriothecium ( x 400). Morenoina arundinariae sp.nov. (Fig. 3) Morenoina chamaecyparidis sp.nov. (Fig. 4) Mycelium abundans, ex hyphis brunneis 2-2-5 Pm latis Mycelium sparsim, ex hyphis pallide brunneis, 1-2 fim compositum. Thyriothecia simplicia vel ramosa, recta latis compositum. Thyriothecia simplicia, dispersa, vel flexa, dispersa vel aggregata, 400-noo x 60-100/im. 120-450 x 80-100 fim. Scutellum ex cellulis atro- Scutellum ad centrum opacum, marginem versus rufo- brunneis, oblongis compositum; cellulae marginales brunneum, cellulae marginales protrudentes. Asci fumoso-brunneae non protrudentes. Asci 14-20 x 25-40 x 13-18 /an. Ascosporae l-septatae, 15-18 x 8-9 fim. Ascosporae l-septatae, 8-10 x 3-4 fim, hya­ 4-6 /im, hyalinae vel stramineae, breviter echinulatae. linae vel pallide brunneae, minute et dispersim verru- Pycnothyria non visa. culosae. Pycnothyria non visa. In caulibus putrescentibus Arundinariae, Plas-Tan- In foliis emortuis Chamaecyparidis lawsonianae, y-Bwlch, Gwynedd, 15 May 1976, J. P. Ellis, Holo­ Hartland Forest, N. Devon, 22 May 1972, J. P. Ellis, typus IMI 204120. Holotypus IMI 168863.

Mycelium abundant, hyphae brown, 2-2-5 /

Fig. 3. Morenoina arundinariae. (A) Habit; (B) parts of edge of scutellum; (C) asci and ascospores. On dead stems of Epilobium angustifoliutn. This Morenoina clarkii sp.nov. (Fig. 5) description is based on Libert's material in Herb.K, Mycelium abundans, ex hyphis brunneis, sinuatis, Plant, crypt. Arduennae: 273 (1834), an isotype 2-4 fim latis compositum, ad hyphas subcuticulars, collection and is included as it may well occur in septatas, torulosas passim affixum. Thyriothecia Britain. simplicia vel ramosa, flexuosa, 250-1500 x 80-120 fim, interdum confluentia. Scutellum ad centrum opacum; cellulae marginales brunneae, saepe in mycelium Morenoina festucae (Lib.) J. P. Ellis comb.nov. crescentes. Asci 25-35x 12-17 f-m. Ascosporae 1- (Fig. 7) septatae, 14-19 x 4-5-6-5 /im, hyalinae vel stramineae Aulographumfestucae Lib., Plant, crypt. Arduennae: ut videtur semper glabrae. Pycnothyria non visa. 373 (1837)- In caulibus emortuis Rubi fruticosi agg., Chesterton Wood, Warwicks., 15 Oct. 1970. M. C. Clark 748, Echidnodes festucae (Lib.) Ryan, III. Biol. Monogr. Holotypus IMI 151898. 17 (2): 97 (1939)- Mycelium plentiful; hyphae brown, 1^5-2-5/*¾ Mycelium abundant; hyphae brown, contorted, wide, often branching at right angles. Thyriotliecia nodulose, 2-4 fim wide, at intervals connected to scattered, simple, bent and variously branched, subcuticular mycelium composed of hyaline, 125-400(800) x 40-80 fim. Scutellum dark red- septate, swollen hyphae. Thyriothecia mostly bent, brown in the centre, paler at the fimbriate margin, lobed and branched, 250-1500 x 80-120 fim, up to with many cells running out into the mycelium. 250 fim wide where lobed, and sometimes con­ Asci 18-22 x 8-ii fim. Ascospores l-septate, 8-9 x fluent. Scutellum almost opaque except at the edge 2-5-3 /*mj hyaline to straw-coloured, minutely where the brown cells frequently run out into the x verruculose. Pycnothyria not seen. mycelium. Asci 25-35 12-17 /*rn. Ascospores l-septate, 14-19 x 4-5-6-5 fim, hyaline to pale On dead sheaths of Festuca sylvatica Vill. (F. straw-coloured, remaining smooth. altissima AIL). This description is based on part of the type collection of Libert in Herb.K. Specimens On dead stems of Rubus fruticosus agg. Five referred to this species as Aulographum festucae collections examined from Devon and Warwicks. have been recorded from this country but none that This fungus is named after Mr and Mrs M. C. I have seen can be placed here. Festuca altissima is a Clark, who have made all the collections. locally rather rare grass of rocky slopes and wooded MORENOINA EPILOBII (Lib.) von Arx, Beitr. stream sides scattered over Europe, and in Britain Kryptogamenfl. Schweiz 11: 129 (1962). mainly in the west and north. (Fig. 6). Morenoina fimbriata sp.nov. (Fig. 8) Mycelium sparse; hyphae straw-coloured to pale brown, somewhat torulose, 1-5-2-5 fim wide. Mycelium moderate abundans, ex hyphis pallide Thyriothecia simple, branched or bent, 150-450 x brunneis 1-5-2 fim latis compositum. Thyriothecia 40-60 fim. Scutellum brown, slightly darker in the simplicia, interdum ramosa, saepe flexa, 300-1100 x 80-120 fim, utrinque angustata. Scutellum ad centrum centre, composed of almost square cells, somewhat atrobrunneum, margine pallidum; cellulae marginales fanned out at the ends, margin unfringed. Asci x saepe in mycelium crescentes. Asci 20-25 12-18 /

10 Mm

Fig. 4. Morenoina chamaecyparidis. (A) habit; (B) edge of scutellum; (C) ascus and ascospores. Fig. 5. Morenoina clarkii. (A) Habit (x 6); (B) thyriothecia (x too); (C) asci (x 1000); (D) edge of scutellum; (E) ascus and ascospores.

In foliis, vaginis et culmis emortuis Junci effusi, 12-18 fan. Ascospores l-septate, 12-15 x 4-5-5 /*m> Caricis binervis et Caricis sp., Hartland Forest, N. hyaline to pale straw-coloured, distantly verru- Devon, 22 May 1972, J. P. Ellis. Holotypus IMI culose. 166599. On dead leaves, sheaths and culms of Juncus Mycelium moderately abundant; hyphae pale effusus, Carex binervis and Carex sp. Four collec­ brown, 1-5-2 fim wide. Thyriothecia scattered, tions examined from Devon, Gwynedd, Powys and simple, sometimes branched, often curved, tapering Warwicks. at each end, 300-1100 x 80-120 fim. Scutellum dark Morenoina minuta sp.nov. (Fig. 9) brown at the centre, paler at the margin, marginal Mycelium primum abundans dein sparsum, ex hyphis cells frequently running out into the mycelium stramineis vel brunneis, 1-5-2 fim latis compositum. giving a fimbriate appearance. Asci 20-25x Thyriothecia simplicia, recta vel interdum flexa, B

Fig. 6. Morenoina epilobii. (A) Thyriothecia ( x 250); (B) ascus and ascospores.

I I 10 /im Fig. 7. Morenoina festucae. (A) Thyriothecia (x 100); (B) edge of scutellum; (C) ascus and ascospores.

100-250 x 40-60 /im. Scutellum rufo-brunneum; cellu­ Mycelium abundant but becoming sparse with lae marginales pallidiores, saepe protrudentes et age; hyphae straw-coloured to brown, 1-5-2//m conicae. Asci 15-20 x 9-12 /im. Ascosporae i-septatae, wide. Thyriothecia straight or curved, unbranched, 7-9 x 2-5-3-5 /"-m> hyalinae vel pallide stramineae, 100-250 x 40-60 fim. Scutellum red-brown, cells interdum minute et dispersim verruculosae. Pycno­ thick-walled, many of those at the margin pro­ thyria non visa. truding and cone-shaped. Asci 15-20 x 9-12 fim. In foliis, vaginis et culmis emortuis Junci effusi, Caricis flaccae, Caricis sp., et Cladii marisci, Cnwc, Ascospores l-septate, 7-9 x 2-5-3-5/im, hyaline to Mynnydd Prescelly, Dyffyd, 25 May 1961, J. T. pale straw-coloured, occasionally minutely and Palmer, Holotypus IMI 134295. distantly verruculose.

Fig. 10. Morenoina paludosa. (A) Habit (x6); (B) edge of scutellum; (C) edge of pycnothyrium; (D) asci and ascospores; (E) conidia.

On dead sheaths, leaves and stems of Juncus Scutellum composed of brown, almost square cells. effusus, Carex flacca, Carex sp. and Cladium Conidia hyaline, cylindrical, 5-7 x 1-5-2 fim. mariscus. Eight collections examined from Cambs., On dead culms, leaves and sheaths of Juncus Devon, Dorset, Dyffyd, Gwynedd, Suffolk and effusus, Carex elata and Cladium mariscus. Nine Warwicks. collections examined from Cambs., Cornwall, As the name suggests, this species is very small Gwynedd, Hunts, and Norfolk. but appears to be quite widespread. The conical Under a lens this is very like the next species cells at the edge of the scutellum seem to be M. phragmitidis, appearing as spots and stars and diagnostic. must be examined microscopically for correct determination. The larger conidia are quite diag­ Morenoina paludosa sp.nov. (Fig. 10) nostic; the ascospores are also slightly longer. On Cladium mariscus the pycnothyria are much larger Mycelium sparsim; ex hyphis hyalinis vel stramineis, than they are on Carex and Juncus. 1-5-2 fim latis compositum. Thyriothecia simplicia vel multiramosa, 160-800 x 60-100 /tm, saepe confluentia Morenoina phragmitidis sp.nov. (Fig. 11) et dein latiora. Scutellum ex cellulis quadratis, atro- brunneis compositum; cellulae marginales pallescentes Mycelium sparsim; ex hyphis olivaceis vel pallide et longiores. Asci 20-28 x 10-13 /*m. Ascosporae 1- brunneis, 1-5-2 fim latis compositum. Thyriothecia septatae, 9-11-5 x 3-4 /im, hyalinae vel stramineae, simplicia vel ramosa, 320-700 x 80-160 fim, saepe saepe minute et dispersim verruculosae. Pycnothyria multiramosa et confluentia et dein ad 700 x 500 fim. orbicularia 60-120 (400) fim diam. Scutellum ex cellulis Scutellum ad centrum opacum, cellulae marginales brunneis, quadratis compositum. Conidia hyalina, elongatae, olivaceo-brunneae, tenuiter divisae. Asci cylindrica, 5-7 x 1-5-2 fim. 20-30 x 10-13 fim. Ascosporae l-septatae, 8-10 x In foliis, vaginis et culmis Junci effusi, Caricis elatae 3-4 fim, hyalinae vel stramineae, minute et dispersim et Cladii marisci, Lambourne Hill, Perranzabuloe, verruculosae. Pycnothyria orbicularia, pallide brurmea, Cornwall, May 1952, M. B. Ellis, Holotypus IMI 60-120 (200) fim diam. Conidia hyalina, cylindrica, 50152. 3-4 x 0-5-1 fim. In culmis emortuis Phragmitidis australis et Phala- ridis arundinaceae, Wheatfen Broad, Norfolk, 26 Aug. Mycelium scanty; hyphae where present hyaline 1970, E. A. Ellis, Holotypus IMI 154593. to straw-coloured, 1-5-2 fim wide. Thyriothecia simple to much branched, 160-800 x 60-100 fim, Mycelium sparse, disappearing with age; hyphae but where confluent and many lobed the area olivaceous to pale brown, 1-5-2 fim wide. Thyrio­ covered is greater. Scutellum composed of deep thecia simple or branched, 320-700 x 80-160 fim, brown almost square cells becoming longer and when multibranched and confluent up to 700 x paler at the edge. Asci 20-28 x 10-13 fim. Asco­ 500 fim. Scutellum opaque in the centre, marginal spores l-septate, 9-11-5 x 3-4 fim, hyaline to pale cells olivaceous brown, elongate and finely divided. brown, often distantly verruculose at maturity. Asci20-30 x 10-13/im. Ascospores l-septate, 8-10 x Pycnothyria orbicular, 60-120(400) jum diam. 3-4 fim, hyaline to straw-coloured, often minutely Fig. li. Morenoina phragmitidis. (A) Habit (x6); (B) parts of thyriothecia (X250); (C) edge of scutellum; (D) ascus and ascospores; (E) conidia.

and distantly verruculose. Pycnothyria orbicular, septate, 8-10 x 3-4 fim, hyaline to pale brown, pale brown, 60-120(200) fim diam. Conidia hya­ smooth to minutely and distantly verruculose. line, cylindrical, 3-4 x 0-5-1 fim. Pycnothyria orbicular, straw-coloured to pale On dead culms of Phragmites australis and Pha- brown, up to 100 fim diam. Conidia hyaline, laris arundinacea. Four collections examined from cylindrical, 2-3 x 0-5-1 fim. Norfolk and Warwicks. On dead twigs of Rhododendron ponticum and Vaccinium myriillus. Four collections examined Morenoina rhododendri sp.nov. (Fig. 12) from Devon, Dorset and the Isle of Mull. Mycelium sparsim, ex hyphis olivaceis vel brunneis, 1-1-5 i"m latis compositum. Thyriothecia simplicia vel ramosa, 300-400 x 60-120 /im, interdum confluentia. Morenoina websteri sp.nov. (Fig. 13) Scutellum ad centrum opacum, cellulae marginales Mycelium ad margines thyriotheciorum abundans, rufo-brunneae,nonprotrudentes./3sa'l6-20 x 8-13/Mn. aliter sparsim, ex hyphis pallide brunneis vel brunneis, Ascosporae i-septatae, 8-10 x 3-4 fim, hyalinae vel 1-5-2 fim latis compositum. Thyriothecia multiramosa, pallide brunneae, minute et dispersim verruculosae. 400-800 x 150-200 /im, saepe confluentia et dein ad Pycnothyria orbicularia, straminea vel pallide brunnea, 800 x 700 fim. Scutellum ex cellulis atrobrunneis, ad 100 fim diam. Conidia hyalina, cylindrica, 2-3 x irregulariter quadrangularis compositum; cellulae 0-5-1 /im. marginales elongatae fimbria formantes. Asci 30-35 x x In ramunculis emortuis Rhododendri pontici et 10-12 fim. Ascosporae i-septatae, 10-14 4-5 fim, Vaccinii myrtilli, Studland, Dorset, 27 May 1961, hyalinae vel stramineae, minute et dispersim verrucu­ K. Pirozynski, Holotypus IMI 89616. losae. Pycnothyria non visa. In culmis emortuis Arundinariae sp., Hengwent Mycelium sparse; hyphae olivaceous to pale Estate, Gwynedd, 15 May 1976, J. Webster, Holo­ brown, 1-1-5 P-m wide. Thyriothecia simple or typus IMI 204122. branched, 300-400 x 60-120 fan, sometimes con­ fluent and covering a large area. Scutellum opaque Mycelium scanty except at the margins of the at the centre, red-brown at the margin, non- thyriothecia where it is abundant; hyphae pale to fimbriate. Asci 16-20 x 8-13 fim. Ascospores 1- mid brown, 1-5-2 fim wide. Thyriothecia much Fig. 12. Morenoina rhododendri. (A) Habit ( x 6); (B) thyriothecium and pycnothyria ( x 150); (C) con­ fluent thyriothecia (x 150); (D) edge of scutellum; (E) asci and ascospores; (F) edge of pycnothyrium; (G) conidia. branched, 400-800 x 150-200 fim, sometimes con­ the Director, Royal Botanic Gardens, Kew, for fluent and then often covering an area up to permission to examine material in Herb.K whilst goo x 700 fim. Scutellum composed of irregularly I was working at the C.M.I. shaped quadrilateral cells running out at the margin into a dense mycelial fringe. Asci 30-35 x 10-12 fim. REFERENCES Ascospores l-septate, 10-14 x 4-5/on, hyaline to pale straw-coloured, minutely and distantly verru­ DESMAZIERES, J. B. H. J. (1843). Dixieme notice sur quelques plantes cryptogames la plupart inedites, culose. recemment decouvertes en France, et qui vont On dead culms of Arundinaria sp. Only one paraitre en nature dans la collection publiee par collection has been seen. Named after Prof. John l'auteur. Annales des Sciences Naturelles, partie Webster. Under a hand-lens this species is seen to Botanique, ser. 2, 19, 362. be composed of star-shaped ascomata whereas in LIBERT, M.-A. (1834). Plantae cryptogamiae Arduennae M. arundinariae they are more robust, elongated exsiccatae 3, 272. and branched. I have examined Aulographum LUTTREIX, E. S. (1973). Loculoascomycetes. In The arundinariae Cooke and it is quite different from Fungi: An Advanced Treatise, vol. IV A (ed. G. C. these two Morenoina species on Arundinaria. Ainsworth, F. K. Sparrow and A. S. Sussman), PP- i35-219- New York, London: Academic Press. MULLER, E. & ARX, J. A. VON (1962). Die Gattungen I wish to thank Mr D. W. Fry and my husband, der didymosporen Pyrenomyceten. Beitrage zur Dr M. B. Ellis, for help with the illustrations, and Kryptogamenflora der Schvjeiz 11, 1-922.

{Received for publication 4 April 1979)