22
[ 18 5] Trans. Br. mycol. Soc. 51 (2), 185-206 (1968) Printed in Great Britain NOTES ON MICROFUNGI. III By K. A. PIROZYNSKI* AND G. MORGAN-JONES Commonwealth Mycological Institute, Kew, Surrey (With 9 Text-figures) Fuller descriptions or explanatory notes are given of 14 species of microfungi. Four new combinations, Ceriospora polygonacearum (Petr.), Cryptosporiopsis nobilis (Sacc.), C. turgida (Berk. & Br.) and Didymellaprominula (Speg.), are made and a new name, Phomajolyana, is proposed. As in the previous papers (Sutton & Pirozynski, 1963, 1965) special attention is paid to British microfungi, but species found elsewhere which are closely rel ated to, or involved in, the history of the native fungi under consideration are included. A number of rare British coelomycetes are re- described and illustrated. Additions to the British fungus flora are marked with an asterisk. Unless otherwise stated, the material or preparations made from the material examined and cited are deposited in Herb. 1M!. BLOXAMIA TR UNCATA Berk. & Br., in Ann. Mag. nat. Hist, I3, 468, 1854. (Fig. I) = Hormococcus nitidulus Sacc., in Michelia, 2, 285, 1881. = Trullula nitidula (Sacc.) Sacc., in Sylloge Fungorum, 3, 732, 1884. = Bloxamia nitidula (Sacc.) Hohnel , in Annls mycol. I, 405, 1903. = Bloxamia saccardiana Allesch., in Rabenhorst, Kryptogamenflora von Deutschland, Oesterreich und der Schioeiz, I, Abt. 7, 553, 1903. The genus Bloxamia Berk. & Br. is based on B. truncata Berk. & Br., a rarely collected tub erculariaceous fungus occurring in this country on dead, decorticated wood of Ulmus. Sporodochia black, at first punctiform, sometimes scattered or more frequently gregarious to confluent, 140-180 JL diam, when confluent, pustule up to 500 JL diam. Mycelium composed of branched, septate, subhyaline to very pale brown, smooth-walled, 2-2'5 JL thick hyphae. Stromata superficial, pale brown, made up of long, loosely aggregated, vertically arranged cells, 120-160 x 25-35 JL. Conidiophores arising termin- ally from the upper cells of the stromata, straight, cylindrical, arranged in a close palisade, subhyaline to pale brown, smooth-walled. Each conidio- phore consists of a septate stipe and a long phialide, 18-35 x 3'4-4'5 JL. Conidia formed endogenously (endophialospores) in basipetal succession (up to six conidia can usually be seen within the spore sac), irregularly cuboid, sometimes oblong, truncate at both ends, guttulate, smooth- walled, hyaline, 2-3 XI. 5- 2 JL. Present address : Plant Research Institute, Central Experimental Farm, Ottawa, Canada.

Notes on microfungi. III

Embed Size (px)

Citation preview

Page 1: Notes on microfungi. III

[ 185 ]

Trans. Br. mycol. Soc. 51 (2), 185-206 (1968)Printed in GreatBritain

NOTES ON MICROFUNGI. III

By K. A. PIROZYNSKI* AND G. MORGAN-JONES

Commonwealth Mycological Institute, Kew, Surrey

(With 9 Text-figures)

Fuller descriptions or explanatory notes are given of 14 species of microfungi.Four new combinations, Ceriospora polygonacearum (Petr.), Cryptosporiopsis nobilis(Sacc.), C. turgida (Berk. & Br .) and Didymellaprominula (Speg.), are made and anew name, Phomajolyana, is proposed.

As in the previous papers (Sutton & Pirozynski, 1963, 1965) specialattention is paid to British microfungi, but species found elsewhere whichare closely related to, or involved in, the history of the native fungi underconsideration are included. A number of rare British coelomycetes are re­described and illustrated. Additions to the British fungus flora are markedwith an asterisk. Unless otherwise stated, the material or preparationsmade from the material examined and cited are deposited in Herb. 1M!.

BLOXAMIA TRUNCATA Berk. & Br., in Ann. Mag. nat. Hist, I3, 468, 1854.(Fig. I)

= Hormococcus nitidulus Sacc., in Michelia, 2, 285, 1881.= Trullula nitidula (Sacc.) Sacc., in Sylloge Fungorum, 3, 732, 1884.= Bloxamia nitidula (Sacc.) Hohnel, in Annls mycol. I , 405, 1903.= Bloxamia saccardiana Allesch., in Rabenhorst, Kryptogamenflora von

Deutschland, Oesterreich und der Schioeiz, I, Abt. 7, 553, 1903.

The genus Bloxamia Berk. & Br. is based on B. truncata Berk. & Br., ararely collected tuberculariaceous fungus occurring in this country ondead, decorticated wood of Ulmus.

Sporodochia black, at first punctiform, sometimes scattered or morefrequently gregarious to confluent, 140-180 JL diam, when confluent,pustule up to 500 JL diam. Mycelium composed of branched, septate,subhyaline to very pale brown, smooth-walled, 2-2'5 JL thick hyphae.Stromata superficial, pale brown, made up of long, loosely aggregated,vertically arranged cells, 120-160 x 25-35 JL. Conidiophores arising termin­ally from the upper cells of the stromata, straight, cylindrical, arranged ina close palisade, subhyaline to pale brown, smooth-walled. Each conidio­phore consists of a septate stipe and a long phialide, 18-35 x 3'4-4'5 JL.Conidia formed endogenously (endophialospores) in basipetal succession(up to six conidia can usually be seen within the spore sac), irregularlycuboid, sometimes oblong, truncate at both ends, guttulate, smooth­walled, hyaline, 2-3 X I . 5-2 JL.

• Present address : Plant Research Institute, Central Experimental Farm, Ottawa,Canada.

Page 2: Notes on microfungi. III

186 Transactions British Mycological Society

Hab, On wood of Ulmus montana, Bath, Somerset, 5 March 1852'C. E. Broome, holotype of Bloxamia truncata (K); Bath,Somerset,February1856 (Rabenhorst, Fungi europaei no. 168), April 1859, April 1865, and

:,:.' . ....

:~ ' -.~ ::

... . ...:: , , -:;.... .

"':. ,::::". ::',

Fig. I. Bloxamia truncata, from type.

Page 3: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-fones 187February 1871 (M. C. Cooke, Fungi Britannici exsiccati no. 472), C. E.Broome (K); on wood ofPyrus malus, Selva, Italy, autumn 1880 (Saccardo,Mycotheca Veneta no. 1598), isotype of Hormococcus nitidulus (IMI 123246ex K).

The mode of conidium ontogeny corresponds closely to that occurringin Sporoschisma Berk, & Br, and Chalara (Corda) Rabenh.

In some details of the conidiophores Bloxamia truncata bears a closeresemblance to Chalara. In C. cladii, Ellis (1961) noted that' the elasticwall around the mouth contracts slightly and seems to clasp the issuingconidia quite firmly'. This is frequently the case in B. truncata. Occasionallywhen a spore sac has ceased to produce conidia, proliferation from the basemay occur through the old sac to produce another sac at a higher level.A similar process was described by Ellis in C. cladii.

The hyaline nature of the conidia and the lack of capitate hyphae inthe conidiophore layer would place B. truncata nearer Chalara thanSporoschisma.

Hohnel (1902) suggested that Bloxamia Berk. & Br., Trullula Ces. andSeptotrullula Hohn, are closely related, and later (1g03) placed Bloxamianear to Hymenula Fr.

Grove (1937) also considered Bloxamia to be a closely related genus toT rullula Ces. The method of conidium production in the two genera is,however, quite different, and furthermore there is little resemblance inthe form of the fructification.

Sutton & Pirozynski (1965) have shown that in Septottullula bacilligeraHohn. , the conidia are formed acropetally, each one originating from theapex of a previous conidium in a simple blastic manner. This evidence,together with that recorded in this paper, shows conidium ontogeny in thethree genera to be quite different.

Cryptosporiopsis turgida (Berk. & Br.) comb.nov. (Fig. 2A)

== Crytosporium turgidum Berk. & Br., in Ann. Mag. Nat. Hist.ri, 129,1881.== Stagonospora turgida (Berk. & Br.) Sacc., in Sylloge Fungorum, 3,

447, 1884.= Cryptosporiumfraxini Rostr., in K. danske Vidensk. Selsk, Skr, 4,38, 1904.

Fructifications scattered, gregarious or frequently confluent, at first im­mersed, then becoming erumpent and rupturing the periderm, discoid,prominent, dark brown to black, 22lJ-500 JL diam. Stromata partly super­ficial, partly immersed, made up of several layers: a layer of long, sub­hyaline to light brown cells at the base gives rise to a layer of thick-walled,light brown cells about 3lJ-40 JL wide, above which is a layer forming thesuperficial discoid portion composed of small isodiametric, subhyalinecells , 25-30 JL wide in the central part. Conidiophores arising terminally in aloose palisade from the cells ofthe upper layer, straight or more frequentlyflexuous, cylindrical, hyaline, smooth-walled with lateral swellings,phialidic, possessing a small collar at the apex, 30-35 x 3-4 JL. After severalconidia have been formed the inner wall of the phialide may proliferatethrough the collar region to form another collar at a higher level on the

Page 4: Notes on microfungi. III

188 Transactions British Mycological Society

same conidiophore. Conidia broadly falcate, slightly more curved towardsthe apex, with an obtuse upper end and a somewhat truncate base,hyaline, guttulate, smooth-walled, formed successively from the openend of the phialides, 20-28 x 4-5 fL.

Fig. 2. A, Cryptosporiopsis turgida. Fructification, conidiophores and conidia, fromIMI 105567;B, C. nobilis, conidiophores and conidia, from Saccardo, Mycotheca italica no. 1563.

Page 5: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 189

Hab. On dead branch ofFraxinus, Ham Park, nr. Dovedale, Derbyshire,April 1964, G. Morgan-Jones (IMI 105567); on twigs ofFraxinus, Rhostry­fan, Caernarvonshire, Wales, 5 June 1966, G. Morgan-Jones and K. A.Pirozynski (IMI I 19422).

C. turgida bears a close resemblance to Cryptosporiopsis malicorticis (CordI.)Nannf. in most respects. It is considered that Cryptosporiopsis provides amore appropriate placing for it.

Cryptosporiopsis nobilis (Sacc.) comb.nov. (Fig. 2B)

= Gloeospororium nobile Sacco in Michelia, 2, 153, 1880.= Cryptocline nobile (Sacc.) Arx, in Verh. K. Akad. Wet. (Ser. 2),51, 109,1957·Cryptocline nohile (Sacc.) Arx possesses long, cylindrical phialides which

produce through the open end a succession of broadly falcate, guttulateconidia with an obtuse upper end and a subtruncate base. The extremetip of the phialides is broader than the part immediately below, thusgiving the appearance ofa small collar. The phialides are borne terminallyand laterally on septate, sparingly branched conidiophores which arisedirectly from the basal stratum of the fructification composed mostly ofrelatively thin-walled elongated cells. The conidiophores frequently haveprominent lateral swellings. As is the case in C. turgida after a number ofconidia have been produced the phialide may proliferate through theopen end. This feature has not been observed in any species ofCryptocline orGloeosporidiella. The detailed morphology of the conidiophores and conidiaof C. nohile reflects a closer affinity to Cryptosporiopsis Bubak and Kabatrather than to Cryptocline Petrak. Arx (1957) considered C. nobile to be atransitional form to Gloeosporidiella. He further considered Cryptocline as awhole to be quite near to Gloeosporidiella but differing in possessing usuallyellipsoid or elongate conidia which arise from distinct rod-shaped conidio­phores. In the same publication Arx stated that Cryptosporiopsis is alsorelated to Cryptocline and that the two are connected by intermediateforms.

The present authors consider that relationship within this group is bestindicated by conidiophore and conidium morphology. The close re­semblance in this respect of C. nobile to C. malicorticis and C. turgida, inspite of the fact that the fructification in the former is not as well developedas is characteristic ofCryptosporiopsis, leads us to consider it better placed inthat genus.

Hab. On leaves ofLaurusnobilis, Castel Gandolfo, Rome, Italy,July 1904(Saccardo, Mycotheca italica no. 1563.) (PAD). A short description ofthe fungus is provided by Arx (1957).

* Ceriospora polygonacearum (Petrak) comb.nov. (Fig. 3)- Amphorulopsis polygonacearum Petrak, in Sydowia, 13, 182, 1959.

In 1964 one of us collected an interesting ascomycete on stems ofPolygonum sachalinense in Scotland. The fructifications resemble veryclosely those of the pycnidial fungus Amphorula polygoni (Ell. & Ev.)

Page 6: Notes on microfungi. III

190 Transactions British Mycological Society

Petrak which occurs on dead stems of Polygonum and Rumex in Europe(including Britain) and North America, and which was recently investi­gated in detail by Petrak (1959). To the list ofsynonyms quoted by Petrakthat of Robillarda discoides Sacco (Bull. Soc. r. Bot. Belge, 28, 98, 1889) canbe added. We have examined the type specimen (Herb. PAD) collectedon Polygonum polymorphum in Siberia, and found it identical with A. polygoni.

Fig. 3. Ceriospora poiygonacearum, from 1M! 1103°1 a.

Associated with A. polygoni on Rumex, Petrak found an ascomycete whichhe assumed was its perfect state. Though the fungus was immaturePetrak proposed a new genus Amphorulopsis for it and described the speciesas A. polygonaceasum Petrak. A few more mature ascospores found were,according to Petrak, up to 4- or 5-septate, and about 20 x 5 ft. We havenot seen the type specimen, but when examining a Swedish collection ofAmphorula poiygoni or Rumex acetosa (A. G. Eliasson, 6. VII. 1927 sub'Kellermania rumicis') we also found an immature sphaeriaceous ascomycete

Page 7: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 191

associated with, and in shape and structure closely resembling, thepycnidial fungus. Except for the absence of multiseptate spores the fungusfits Petrak's description of Amphorulopsis polygonacearum exactly. The asco­spores are fusiform, slightly curved, hyaline and multiguttulate, with thenarrow strips of cytoplasm separating the guttules strongly reminiscent ofsepta. However, one perithecium examined, though still immature, hadascospores more fully developed: they were one-septate in the middle andbore apical appendages typical for the genus Ceriospora Niess!' In this andall other respects this fungus closely resembles our specimen from Scotland.Although the ascospores are smaller than the fully mature spores from theScottish material, they are identical with the younger spores from ourcollection. We, therefore, consider the two fungi to be, without doubt,conspecific. Furthermore, we are of the opinion that A. polygonacearumalso belongs to the same species. Amphorulopsis is therefore a synonym ofCeriospora and we propose to emend its description accordingly.

Superficially the fungus appears as blackened and slightly bulging areasof epidermis perforated by somewhat raised, papillate ostioles. Thisblackening is due to the presence of a plectenchymatous clypeus composedof dark brown hyphae filling the cells of epidermis. Underneath andusually resting on a xylem strand are black, sclerotium-like stromata,rounded or oval, flattened dorsa-ventrally, simple or compound, the latteroften confluent in a linear manner. These stromata are firmly attached to theepidermis and remain so when the epidermis is loosened and eventuallypeels off old stems. They are 30(}-400 p. diam and 30(}-300 fL deep whensimple, up to 800 p. diam and 400 p. deep when confluent, composed ofbrown parenchyma ofmore or less angular cells ± lOP. across. Two or threerows of outermost cells have thicker walls and are darker in colour.Perithecia flask-shaped, ISO-2S0 p. diam and 12(}-200 p. deep, embedded inthe centre of the stroma. Necks IO(}-150 p. long, lined with conspicuousperiphyses, growing vertically towards the surface where ostioles, eachsurrounded by a collar of elongated cells, pierce the epidermis. Perithecialwall thin, composed of pale yellowish brown, tangentially flattened cells.Asci cylindric to subclavate, arising in a more or less flat layer, not liberatedinto perithecial cavity, briefly pedicellate, 8-spored, 75-9S x la-IS p..Paraphyses hyaline, occasionally septate, 12S-150 p. long, varying in thick­ness from 2'S to YS p.. Ascospores hyaline, more or less regularly distichous,narrowly ellipsoid, r-septate in the middle, 14-28 x 3-6 p. (mostly18-26 x 4'5-YS p.). When mature (but not old) the spores are furnishedwith apical appendages which are flexuous or quite straight and veryvariable in length (6-40 p. long), 1'5-2 p. wide at the base, tapering toan acutely pointed apex. The ascospores are at first small, 14-20 x 3'5 p.,straight or curved, aseptate and strongly guttulate, usually with 4-6 largeguttules. The appendages are not obvious at this stage. Later, while stillin asci or just after liberation the ascospores elongate (sometimes up to32 p.) but remain narrow, 4-S p. wide. They become curved, develop amedian septum but remain unconstricted, and produce clearly visibleappendages. Old ascospores often lose their appendages, become fatter andshorter, little or not curved, constricted at the septum, and the upper celloften becomes shorter and wider, sometimes as wide as 8·S fL.

Page 8: Notes on microfungi. III

192 Transactions British Mycological Society

Hab. On dead stems of Polygonum cuspidatum, Heythrop Park, Oxon.,26 June 1931, W. B. Grove (IMI 124602 ex K); on P. sachalinense Kirk­patrick, Dumfriesshire, Scotland, 5 July 1964, K. A. Pirozynski (IMII I030ra); on Rumex acetosa, Malen, Scania, Sweden, 6 July 1927, A. G.Eliasson (IMI 16634b).

Grove (1935) noted that Amphorula polygoni was often accompanied andfollowed by a fungus: which he named as "Diaporthe maculosa j. polygoniGrove'. As far as we can ascertain this name has never been validlypublished and no description exists. We have examined 12 collections ofAmphorula made by Grove on Polygonum sachalinense and P. cuspidatum inSutton Coldfield, Heythrop Park and Edgbaston (Herb. K), and foundexcellent material of Amphorulopsis. Since no other fungus is present weassume that this must be the fungus Grove referred to as 'D. maculosa j.polygoni'. We have also examined four specimens (Herb. BM) collected byGrove on Rumex and labelled' Diaporthemaculosa S. & S'. These turned outto be mixed collections of either Diaporthe arctii (Lasch) Nits. or Plagiostomadeuexa (Desm.) FuckeL

The excellent material of both Amphorula and Ceriospora collected byGrove on P. cuspidatum in Heythrop Park clarified the fact that the fungusis stromatic: conflicting statements exist in the literature concerning thestructure of the fructification. The specimen also yielded valuable informa­tion concerning relationship between the two fungi. A single stroma wasfound containing both a perithecium and a conidiallocule. This, in ouropinion, very strongly supports Petrak's assumption (and our earlierobservations) that the two fungi represent states of the same organism.

Amphorulapolygoni has been adequately dealt with by Petrak (1959), buta word on conidium ontogeny may be of interest.

The conidia are phialospores produced singly and successively from theopen end of long, cylindrical phialides. After a number of conidia havebeen formed at the apex, the conidiophore may develop a new lateralgrowing point. This results in the displacement of the original apex to oneside and a new opening is formed at a higher level through which abasipetal succession of conidia is produced (Fig. 4).

Lind (1913) listed Kellermania rumicis Fautr. & Lamb. (= Amphorulapolygoni (ElL & Ev.) Petrak) as a synonym of Heteropatella cercosperma(Rostrup) Lind (basionym: Septaria cercosperma Rostr.), Rhabdospora caudata(Karst.) Sacco (basionym: Septaria caudata Karst.) was also considered byLind to be the same species. If S. cercosperma Rostr. (1883) or S. caudataKarst. (1884) are the same as K. rumicis either could provide an earlierepithet for Amphorula polygoni (basionym: Kellermania polygoni Ell. & Ev.,1886).

Hohnel (1915) and Petrak (1959) mentioned Lind's work, but neitherauthor endeavoured to clarify the position. S. cercosperma can be disregardedas a possible synonym of Amphorula. We have examined several collectionsdetermined by E. Rostrup as C. cercosperma (Herb. GRO) and find themto be Heteropatella. S. caudata has neither been issued in Karsten's exsiccatanor is there any material of this species in Herb. H (Dr H. Roivainen,personal communication). According to the description, and in Saccardo'sopinion (1892), S. caudata is hardly different from S. cercosperma, Unless

Page 9: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 193Karsten's type is found and shown to be the same as A. polygoni (whichseems unlikely) the name A. polygoni as emended by Petrak must stand.Lind (1926) referred Heteropatella cercosperma to H. umbilicata (Fr.) Jaap,and gave a long list of synonyms which again included Kellermania rumicis.

Fig. 4. Amphorulapolygoni, from IMI 107950.

H. cercosperma had previously been treated by Vestegren (1900) (asRhabdospora centrosperma) , and J aap (1907) (as Heteropatella umbilicata).They did not, however, list Rumex as a host. Collections on Rumex on whichLind based his new combination (1913) are in all probability A. polygoniwhich he misdetermined as Heteropatella.

13 Myc.51

Page 10: Notes on microfungi. III

194 Transactions British Mycological SocietyThe genus Ceriospora and its seggregate Ceriosporella Berlese, 1891 (non

Cavaliere, 1966) was recently investigated by MUller & Arx (1962) whoremoved Ceriosporella from Sphaeriales into Pseudosphaeriales (Lophiosto­mataceae). Ceriospora, as interpreted in its original pre-Winterian sense wasretained in the Sphaeriales. Muller & Arx dealt with two species of Cerio­sporella: C. patouillardii Let., the type species of Ceriosporella, was consideredsynonymous with Ceriospora ulicis Pat. which in turn is a Lophiosphaera(L. ulicis (Pat.) Muller). The second species, C. bicalcarata (Ces.) Berl. wasreverted to its original position in Ceriospora. The remaining three speciesof Ceriosporella listed, namely C. acuta (Smith) Sacc., C. gallica Sacco andC. polygoni Smith & Ramsb. were not investigated.

We have examined the type material of C. polygoni (Herb. BM) andfound it identical with Plagiostoma deuexa (Desm.) Fuckel (see Fig. 8D andDennis, 1960). Diaporthe (Tetrastega) sachalinensis Sacco as exemplified by thetype specimen collected by Fautrey in Cote d'Or and deposited inSaccardo's herbarium (PAD) is the same species.

The type specimen of C. acuta collected by W. R. Elliott in Dominica(Herb. BM) is in very poor condition, but there can be little doubt that itis a Lasiosphaeria.

We have also examined the type collection of Sphaeria curvicolla Peck(Gnomoniella curvicolla (Peck) Sacc.) described from Polygonum articulatumfrom North America and deposited in Herb. NYS. This fungus too is invery poor condition and is a member of Loculoascomycetes, possibly aBottyosphaeria.

In the course of our investigation of Ceriospora on Polygonum and Rumexthree species of Didymella Sacco were examined and are described below.

DIDYMELLA SACHALINENSIS Sacc., in Annls mycol. 6, 558, 1908. (Fig. 5B)

Pseudothecia scattered singly, sometimes gregarious or even confluent,subepidermal, embedded in much disintegrated host tissues which arelargely replaced by a loose weft of dark brown hyphae in the immediatevicinity of the fructification, subglobose, 280-350 I-'- diam, 240-340 I-'- deep,up to 550 I-'- diam when confluent, with a wide and often somewhatpapillate ostiole breaking to the surface. Pseudothecial wall 20-35 I-'- thickcomposed of cells 5-8 I-'- across, rounded, dark brown and rather obscureon the outside, pale yellow brown to subhyaline and progressively moreangular and tangentially flattened towards the centre. Asci arranged ina more or less flat layer, numerous, narrowly cylindrical, shortly pedi­cellate, 8-spored, 110-140, sometimes up to 170 I-'- long, 10-13 I-'- wide.Pseudoparaphyses numerous, filiform, septate, occasionally branched.Ascospores monostichous, very rarely distichous and then only in the upperpart, lying obliquely in an ascus, widely spaced, rarely overlapping,broadly ellipsoid to ovoid, hyaline, one-septate in the middle and moder­ately constricted at the septum with the upper cell slightly wider, 15-17 x5·5 I-'- when in asci, becoming up to 20 I-'-longand 91-'- wide after liberation.

Hab. On dead stems ofPolygonum sachalinense, Rigny sur Arroux, France,1908, J. A. Flageolet (IMI 120927 ex PAD), holotype,

Page 11: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 195

10p I

A 1.-1 _~_--J

Fig. 5. A, Didymella prominula (a-e, ascospores from different collections, see text);B, D . sachalinensis, from type.

Page 12: Notes on microfungi. III

196 Transactions British Mycological Society

LOPHIOSPHAERA FUCKELII Sacc., in Michelia, I (3), 336, 1878. (Fig. 8E)

= Didymosphaeria (Didymella) lophospora Sacco & Speg. ap. Sacc., inMichelia, I (4), 376, 1878; Fungi ital. tab. 367.

= Didymella lophospora (Sacc. & Speg.) Sacc., in Sylloge Fungorum, I, 561,1882.

= Didymella lophospora (Sacc. & Speg.) Sacco var. acetosellae Ellis & Saccoap. Sacc., in Michelia, 2, 374, 1881.

= Lophiotrema lophosporum (Sacc. & Speg.) Rehm, in Annls mycol. 5,5 I 8, 1907 (as 'lophiosporum ').

Rehm (1907) published the new combination Lophiotrema 'lophiosporum'(Sacc. & Speg.) Rehm, after examining Ellis's exsiccatum (N. Am. Fungino. 588) labelled 'Sphaeria (Didymella) lophospora Sac. & Speg.'. We haveexamined the portion of N. Am. Fungi no. 588 in Herb. K. This is amixed collection containing (a) dead stems of Rumex acetosella and (b) oldcapsules of Oenothera. The part (a) on Rumex is almost certainly the type ofD. lophospora var, acetosellae, and we agree with Rehm that the fungusbelongs to the Lophiostomataceae. It is, in fact, a species of LophiosphaeraTrev. which in our opinion does not sufficiently differ from L.fuckelii Saccoto warrant separation from this species. We have also examined an authenticcollection of Didymella lophospora from Saccardo's herbarium (PAD). Thetype specimen is lost and the collection examined is the only specimen ofthe species authenticated by Saccardo available. It was collected by].]. Therry in May 1861 in the Isere region of France (Cryptogames duLyonnais no. 5770) in very small quantity and poor condition. However,a few pseudothecia present have characteristically compressed ostioles,and the asci and ascospores, which incidentally were illustrated bySaccardo on the packet, are those of Lophiosphaera fuckelii Sacco We, there­fore, refer both D. lophospora and D. lophospora var. acetosellae to L.fuckelii.

Petrak (1940) recorded Didymella lophospora from stalks of bracken inAustria. He did not observe appendaged ascospores which he explainedwas due to over-ripeness of the fructifications.

Petrak (1963) again recorded the fungus on the fronds of the same hostin Austria (Mycotheca generalis no. 233) (Fig. 5A, a) and suggested itrepresented the perfect state of Ascochyta pteridis Bres. This fungus, deter­mined and recorded (Corbaz, 1957b) as D. lophospora, was also collectedby E. MUller & R. Corbaz on stalks of bracken in southern France (Herb.K) (Fig. 6A, b). We have examined the two collections last mentioned andfound them identical with Sphaerella prominula Speg. The type specimencollected by Spegazzini on bracken in Conegliano, northern Italy, in May1878 is not available. We have, however, been able to see a specimencollected by Spegazzini on the same host and in the same locality in thespring of 1879 and issued in the form of exsiccati underSphaerellaprominula(C. Spegazzini-Decades Myc. Ital. no. 44) (Herb. K, PAD) (Fig. 5A, c).We are of the opinion that this fungus should be placed in Didymella andpropose to amend the description accordingly.

Page 13: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones

501'

Fig. 6. Mycosphaerella limbaIis, from IMI 119036.

197

Page 14: Notes on microfungi. III

Transactions British Mycological Society

Didymella prominula (Speg.) comb.nov. (Fig. 5A)

== Sphaerella prominula Speg., in Michelia, :I, 456, 1879.== Mycosphaerella prominula (Speg.) Lindau, in Engler & Prantl, Die

natiirlichen Pfionzerfamilien, I, Abt. I, 426, 1897.

Fructifications occurring on stems and fronds of bracken. When on stemsgregarious and maculiform, on fronds scattered, epiphyllous. Pseudotheciaimmersed, covered by bulging out epidermis; rounded to conical, 120­180 J.L diam, 80-140 J.L deep, opening by an apical pore. Pseudothecialwall two to four layers of cells thick, composed of olive brown cells whichare somewhat flattened tangentially and ± 10 J.L across, The cells are oftendarker towards the apical pore. Asci not fasciculate but arranged in amore or less flat layer, numerous, centrally placed asci straight, peripheralasci curved to fit the shape of the pseudothecial venter. They are subclavateto cylindric, almost sessile, 8-spored, 50-75 x 10-16 J.L.

Pseudoparaphyses filiform, numerous in young fructifications, not observedin more mature ones. Ascospores irregularly distichous, ovoid to ellipsoid,usually somewhat curved, hyaline, one septate in the middle and stronglyconstricted at the septum, with the upper cell often slightly wider, at first13'5-19 x 4'5-7 J.L, later 18'5-20 x 6'5-9 J.L when fully mature.

Stated by Spegazzini to be very rare, this species is now known fromItaly, Austria, France and Denmark (Munk, 1957) and probably alsooccurs in Britain where, however, it is yet to be collected.

*MYCOSPHAERELLA LIMBALIS (Pers.) Arx, in Sydowia3,87, 1949. (Fig. 6)

Hab. On living leaves of Buxus semperoirens, Oxton House, nr. Dawlish,Devon, 2 May 1966, Miss C. Booth (IMI 119036).

This fungus was first described by Persoon as Phyllosticta limbalis froman immature collection made by Morthier on box leaves near Neuchatel,Switzerland. Arx (1949) reported it to be widespread near Oberbuchsiten,Switzerland. However, judging by the few collections of it made theorganism would appear to be rather rare. Furthermore, it is seldom repre­sented in herbaria in a ripe stage (E. Muller, personal communication).

Arx (1949) described the ascoma wall as being rather dark around theostiole but otherwise light, usually yellow-brown. Because of this heconsidered that M. limbalis could probably be placed in a monotypicsection apart from Eu-Mycosphaerella in which it would otherwise fit.

Mycosphaerella is not the best genus in which to accommodate thisspecies. It can equally well be placed in Didymella with which it has manycharacters in common. It is, however, beyond the scope of this paper todiscuss generic concepts and review the voluminous literature concerningthese two genera. We therefore exercise a conservative attitude to possiblename changes until generic distinctions are clarified and maintain M.limbalis and D. sachalinense for the time being. In transferring M. prominulaas above to Didymella we are perpetuating for it the generic name in whichit is known and under which, in our opinion, following the proposals ofCorbaz (1957a), it is best accommodated.

Page 15: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 199

MYXORMIA ATROVIRIDIS Berk. & Br., in Ann. Mag. nat. Hist., 5, 457,185°.(Fig. 7)

== Crocicreas atroviride (Berk. & Br.) Hohnel, in Annls mycol. 1,4°3, 1903.

Fructifications usually scattered, discoid , superficial, up to 800 fL diam.Mycelium immersed in the substratum, composed of brown, branched,septate, smooth-walled hyphae, 3"5 fL wide. The base of the fructification

Fig. 7. My;connia atroviridis, from IMI 124771.

is lined by a narrow layer ofbrown cells and this gives rise at the sides to anextended exciple, 15-20 fL at the base, narrowing upwards, composed oflong, parallel-walled, olive-green cells. A layer of relatively thin-walled,subhyaline cells, 18-20 fL high, lies within this and gives rise to a loose

Page 16: Notes on microfungi. III

200 Transactions British Mycological Society

palisade of conidiophores. Conidiophores straight or slightly flexuous,cylindrical, branched, septate, hyaline, phialidic, tapering very slightly atthe apex, 20-40 x 1·5 {-to Conidia formed successively through the open endsof the phialides, fusiform to elliptical, sometimes somewhat cylindrical,hyaline to very light olivaceous, smooth-walled, 9-12 x 1.5-2 {-to

Hab. On dead leaves ofDeschampsia caespitosa, Bath, Somerset, December1858, C. E. Broome (K); on D. caespitosa, Bath, Somerset, January 1859,C. E. Broome (a number of duplicates including one issued by Raben­horst, Fungi europaei no. 63) (IM1 124771 ex K); on dead grass, Saunder­ton, Bucks., IS July 1945, R. W. G. Dennis (K); on grass, near Hatten­heim, Germany (Fuckel, Fungi Rhenani no. 548 sub Crocicreas gramineum(Fr.) Fr.).

In addition, we have examined collections sub Perisporium gramineumFr. ex herb. Montagne (Herb. K) on Phleum nodosum collected in theArdennes but have failed to find fructifications on them.

The conidia in Fuckel's collection are consistently cylindrical andnarrower, 1'5 {-t, than those in Broome's collection which are 2 (-t wide.We do not consider this to be ofsufficient significance to prevent placing theformer collection in M. atroviridis.

Petrak & Sydow (1923) reported examining the material on whichFries (1829) based the name Perisporium gramineum and later erected thegenus Crocicreas (Fries, 1849). They consider the collection to be a discomy­cete, possibly a graminicolous species of Phialea, in an immature state.They recommended that the generic name Crocicreas be dropped. Sub­sequently Petrak & Sydow (1937) reiterated this view. Accordingly weadopt the generic name Myxormia Berk. & Br. for the fungus describedabove.

In his preamble to the genus Bloxamia Berk, & Br., Grove (1937)stated that it somewhat resembles Crocicreas Fr. (as Crocicreas Hohnel). Hepresumably based his comment on his knowledge of M. atroviridis. Ourstudies fail to confirm this-there is little ifany resemblance between them.

Phoma jolyana nom.nov,

= Peyronellaea musae Joly, in Revue Mycol. 26, 97, 1961.= Phoma musae (loly) Boerema, Dorenbosch & Kesteren, in Persoonia,

4, 63, 1965. [non Phoma musae Sacc., in Sylloge Fungorum, 3, 163,1884] .

= Peyronellaea nainensis Tandon & Bilgrami, in Curro Sci. 30, 344, 1961.

A new name is proposed for P. musae (loly) Boerema, Dorenbosch &Kesteren since it is a later homonym ofP. musae Sacco Saccardo's name wasin effect a nom.nov. for Sphaeropsis musarum Cooke (in spite of the fact thathis publication of it took the form ofa new combination), necessitated bythe preoccupation ofthe epithet musarum in the genus Phoma by P. musarumCooke. The epithet nainensis employed for the organism under considera­tion by Tandon & Bilgrami cannot be used for the purpose of a transferinto Phoma for the resultant binomial already exists; Phoma nainensisBilgrami (as 'nainense') in Curro Sci. 32, 175, 1963.

The name Phoma musae Carpenter, in Rep. Hawaii agric. Exp. Sin, 1918,

Page 17: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 201

A

1001'

c

[~

Fig. 8. A, Phoma suaedae, from 1M1118388; B, Pseudomassaria chondrospora (a, b, ascosporesfrom different collections, see text); C, Pteridospora tiliae, from IMI 124928; D, Cerio­sporella polygoni, from type (= Plagiostoma devexa); E, Didymellalophospora var, acetosellae,from type (= Lophiosphaera juckelii).

Page 18: Notes on microfungi. III

202 Transactions British Mycological Society

p. 39, erected for the causal organism of banana freckle disease in Hawaiiis an additional later homonym of Phoma musae Sacco

PHOMA SUAEDAE Jaap, in Schr. naturw. Ver. Schlesui-Holst. 14, 27, 1907.(Fig.8A)

Pycnidia gregarious or scattered, at first completely immersed, laterbecoming erumpent, subglobose, 150-23° fJ- diam. The pycnidial wall isusually many cells thick, composed of two layers, an outer layer of darkpseudoparenchymatous thick-walled cells, somewhat darker around theostiole, and an inner hyaline layer bearing conidiophores. Conidiophoresshort, 5-8 fJ- long, hyaline, relatively undifferentiated from the inner cellsof the pycnidial wall, flask-shaped, phialidic. Conidia unicellular, hyaline,elliptical, 5-10 x 3-4 fJ-, biguttulate, formed in basipetal succession asacrophialospores.

Hab. On dead stems of Suaeda maritima, Llandysilio, Menai Bridge,Anglesey, 3 August 1927, P. G. M. Rhodes 2145A (IMI 124753 ex K):Gibraltar Point, Lines., 15 February 1963, G. Morgan-Jones (IMI 118388).

In culture on 2 %malt agar it grows rapidly, producing dense, floccose,white mycelium, and sporulates readily. Immersed mycelium is composedof light brown, smooth-walled hyphae, with the area next to each septumdistinctly swollen to give a bulbous appearance.

It is easily distinguished from other Phoma species on coastal plants byspore size.

Our observations on conidium ontogeny confirm the remarks made bySutton (1964) for other species of the genus.

*PSEUDOMASSARIA CHONDROSPORA (Ces.) Jacz., in Bull. Herb. Boissier, 2,663, 1894. (Fig. 8B)

Hab. On dead twigs of Tilia sp., Slindon, West Sussex, 16 April 1966,D. A. and D. G. Reid (IMI 124928).

The account of the species with extensive synonymy quoted was givenby Muller & Arx (1962) and more recently by Barr (1964).

The fungus occurs in central Europe and southern United States, and ishere recorded from Britain for the first time.

As pointed out by Barr the ascospores show considerable variability inshape and size though both she and Muller & Arx placed them within the17-35 x 7.5-14.5 fJ- range. In Petrak's exsiccati (Mycotheca generalisnos. 350 and 600) which we have examined, the ascospores range fromovoid in shape and approximately 19 x 10 fJ- to narrowly ellipsoid, 30 x 8 JL.According to Barr and Muller & Arx the ascospores are hyaline to yellowishwith guttulate or coarsely granular contents. Petrak's specimens confirmthis (Fig. 8B, b). However, in the British material considerable variabilitywas encountered particularly regarding pigmentation of the ascospores(see also Munk, 1957). The ascospores, apart from quite immaturespecimens which are long and narrow (about 24 x 6 JL), are fairly uniformin size and shape. They are ovoid and measure 18-23 x 8.5-10 JL. When inasci or just after liberation they are hyaline or yellowish but soon become

Page 19: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 203

coloured. The smaller, lower cell remains hyaline or becomes diluteolivaceous but the larger cell soon turns dark brown. Furthermore, theupper cell not uncommonly becomes subdivided by a transverse septumand, in some instances, by two septa (Fig. 8B, a). In spite ofa fair propor­tion of the ascospores obtained from the British collection being stronglypigmented we consider them atypical with both colour and septationbeing due to the age of the collection. Mature, viable ascospores at thetime of liberation from an ascus are colourless or almost colourless. Theygerminate readily on agar media and produce a germ tube up to 1 mmlong overnight. We have encountered some difficulty in growing thefungus beyond germination stage on standard media. Those that grewproduced copious greyish white mycelium which, however, has remainedsterile.

On the same collection we found pseudothecia of a fungus previouslyrecorded from Britain as Massariella curreyi (Tul.) Sacco (Bisby & Mason,1940; Dennis, 1960). This fungus has rightly been transferred by Muller& Arx (1962) into Pteridospora Penz, & Sacco as P. curreyi (Tul.) Muller,who provided an illustration, detailed description and a list of synonymsto which Phorcys tiliae (Curr.) Schroeter (in Cohn, Kryptogamenflora vonSchlesien, 381, 1893-1908) can be added. The ascospores are sparselyminutely roughened and are enveloped in mucilaginous sheath (Fig. 8e).This mucilaginous envelope is of variable thickness and shape and ,oftentakes the form of the space available in the ascus. The first and the lastspore in an ascus often have their sheaths drawn out into tail-like pro­jections.

As pointed out by Bisby & Mason (1940) Massaria tiliae Phill. & Plowr.was considered by Berlese (1894) to be the same as P. eurreyi. However,BisbySc Mason accepted the species and recorded it under Massarina intowhich genus it was transferred by Saccardo (1883). In view of the con­fusion surrounding its position the species should be investigated and theinvestigation must be based on the original material, which we have notbeen able to locate. Unless the type specimen is found the name is bestleft in abeyance.

*SCIRRHIA ASPlDIORUM (Lib.) Bubak, in Ber. At. bot. Ges. 34, 382, 1916.

A full description and illustrations of this species as well as notes onrelated genera and species are provided by Obrist (1959). To the list ofsynonyms quoted that ofSphaeria pteridieola Berk. & Curt., Grevillea, 4, 145,1876 [Didymella pteridieola (Berk. & Curt.) Sacc., Sylloge Fungorum, I, 561,1882] as exemplified by the type specimen (Herb. K), must be added.

Hab. On dead stalks of Pteridium aquilinum, Ranmore Common, Surrey,28 March 1966, G. Morgan-Jones and K. A. Pirozynski (IMI 119041).

Page 20: Notes on microfungi. III

2°4 Transactions British Mycological Society

TRULLULA OLIVASCENS (Sacc.) Sacc., in Sylloge Fungorum, 3, 731, 1884.(Fig. 9)

== Hormococcus olivascens Sacc., in Michelia, 1,94, 1877, and in Fungi ital.tab. 91.

Fructifications scattered, less often gregarious. Conidiophores and conidiaat first enclosed in a subglobose or flattened discoid fructification, 230-300 ILwide, 175-200 IL high. Basal stroma composed ofsmall, isodiametric, light­brown to subhyaline cells. It is covered at the sides and above by an outer

L-...J5/1

Fig. 9. Trullula olivascens, from Saccardo, Mycotheca Veneta no. 1599.

layer of thick-walled, dark brown, loosely aggregated hyphae. Fructifica­tions at first immersed then becoming erumpent and rupturing the cuticleand epidermis. Conidial mass exposed by splitting of the upper part of thefructification which finally becomes excipuloid. Conidiophores in a closepalisade arising from the base and sides of the fructification, long, cylin­drical, hyaline, septate, straight or flexuous, 18-47 x 1.5-2 IL, frequentlybranched, giving rise to long chains of conidia which are at first almostindistinguishable from the conidiophores. Conidia regularly oblong, dis­tinctly truncate at the ends, faintly coloured, olivaceous in mass, smooth­walled, guttulate, 3-6.5 x 1.5-2.5 IL.

Page 21: Notes on microfungi. III

Microfungi III. K. A. Pirozynski and G. Morgan-Jones 205

Hab, On cone scales of Abies sp., Padua, Italy, February 1881, G.Bizzozzero (Saccardo, Mycotheca Veneta no. 1599) (IMI 124616 ex K).

Material of C. Roumeguere's Fungi gallici exsiccati, no. 3186 subHormococcus olivascens Sacc., on pods of Robiniapseudoacacia (Herb. K). doesnot bear the fungus. Petrak's Fungi polonici exsiccati, no. 650, subT rullula olivascens Sacc., on twigs of Berberis vulgaris is apparently a mis­determination. The collection bears a pycnidial fungus with goldenbrown, oval, verruculose conidia associated with a species of Pleospora,

In Trullu/a olioascens the conidiophores develop transverse septa atregular intervals in basipetal succession. The conidiophores break up at thesepta, which presumably attain a double structure to form the oblongconidia with truncate ends. There is little or no thickening of the lateralwalls during this process. At release the wall at the ends can be seen to benoticeably thinner than the side wall of the conidium. A minute frillencircles the thin-walled truncate ends. The double structure of thetransverse septum separates into two parts in the process of differentiationof one conidium from its neighbour. Prior to the breaking' up of the lateralwall a space occurs between the two parts. The break in the lateral wall,which finally causes a conidium to secede from the conidium initial below,occurs at a point between the two close transverse septa which originatedfrom the double structure of the original septum. In this way a smallportion of the lateral wall is carried away at each end ofa given conidium,outside its end walls, as a small frill.

The development of conidia in T. olivascens corresponds closely to thatoccurring in the genus Geotrichum Link ex Fr. representing Hughes' sectionVII (1953).

We wish to thank Prof. C. Cappelletti (PAD), Mrs Balfour-Browne(8M), Drs R. W. G. Dennis (K) and S.J. Smith (NYS) for makingcollections in their keeping available for examination.

REFERENCES

ARx,J. A. VON (1957). Revision der zu Gloeosporium gestellten Pilze. Verh, K. Akad. Wet.(Ser. 2.) 51, 1-153.

BARR, M . E . ( 1964). The genus Pseudomassaria in North America. Mycologia 56, 841-862.BERLESE, A. N. (1894). Icones Fungorum 1, 1-243.BISBY, G. R . & MAsON, E . W. (1940). List of pyrenomycetes recorded from Britain.

Trans. Br. mycol. Soc. 24 , 127-243.CORBAZ, R. ( 1957a) . Phytopath. Z. 28, 375-381.CORBAZ, R. (1957b). Recherches sur Ie genre Didymella Sacco II. Recherches sur

quelques especes de Didymella. Phytopath. Z. 28, 382-414.DENNIS, R. W. G. (1960). British cup fungi and their allies, 1-280. London: Ray Society.ELLIS, M . B. (1961) . Dematiaceous hyphomycctes. II. Mycol . Pap. 79, 1-23.FRIES, E. ( 1829) . Sysuma Mycologicum, pp. 1-269.FRIES, E. (1849) ' Summa Vegetabilium Scandinaviae, pp. 261-572.GROVE, W. B. (1935). British sum- and leaf-fungi (Coelomyceus), vol. I, 1-488, Cambridge

University Press.GROVE, W. B. (1937). British sum- and leaf-fungi (Coelo"!Yceus), vol. II, 1-406, Cambridge

University Press.HOHNEL, F. VON (1902). Fragments zur Mykologie 36. Sitzb , Akad. Wiss. Wien lU, I025.HOHNEL, F. VON (1903). Mycologische Fragments. Annls mycol. 1,391-414.

Page 22: Notes on microfungi. III

206 Transactions British Mycological SocietyHOHNEL, F. VON (1915). Fragments zur Mykologie 900. Uber die Gattung Kellermania

Ellis et Everhart. Sitzb, Akad. Wiss. Wien (Math. -nat.), 124,82-84.HUGHES, S.]. (1953). Conidia, conidiophores and classification, Can.]. Bot. 31, 577-659.JAAP, O. (1907). Beitrage zu Pilzflora der Schweiz. Annls mycoi. 5, 246-272.KARsTEN, P. A. (1884). Fragmenta mycologica. XI. Hedwigia 23,38.LIND,J. (19 I3). Danish fungi, pp. I-648. Copenhagen: Egmont H. Petersen.LIND,J. (1926). Micromycetes from north-western Greenland found on plants collected

during the Jubilee Expedition 1920-23. Meddr Grenland 71, 161-179.MULLER, E. & VON ARx, J. A. (1962). Die Gattungen der didymosporen Pyrenomyceten.

Beitr. KryptogFiora Schuieiz II, 1-g22.MUNK, A. (1957). Danish pyrenomycetes. Dansk. bot. Ark. 17, 1-491.OBRIST, W. (1959). Untersuchungen tiber einige 'dothideale' Gattungen. Phytopath. Z.

35, 357-388.PETRAK, F. (1940). Beitrage zur Kenntnis der Pilzflora der Umgebung von Lunz am See

und des Diirrensteins im Niederdonau. Annis mycol, 38, 121-180.PETRAK, F. (1959). Uber die Gattung AmphorulaGrove und die zu ihr gehorige Schlauch­

form. Sydowia 13, 178-182.PETRAK, F. (1963). Mykologische Beitrage zur osterreichischen Flora. Sydowia 16, 155­

198•PETRAK, F. & SYDOW, H. (1923)' Kritisch-systematische Originaluntersuchungen tiber

Pyrenomyzeten, Sphaeropsideen und Melanconieen. Annis mycoi. 21, 349-384.PETRAK, F. & SYDOW, H. (1937). Uber die Gattung Amerosporium Speg. und ihre

nachsten Verwandten. Annis mycol. 35, 332-338.ROST~UP, E. (1883). Mykologiske Notitser fra en Rejse i Sverige i Sommeren 1882.

(jfvers. K. VetenskAkad. Forh, 4, 35-47.SACCARDO, P. A. (1883)' Sylloge Fungorum 2.

SACCARDO, P. A. (1892). Sylloge Fungorum 10.

SUTTON, B. C. (1964). Phoma and related genera. Trans. Br. mycol. Soc. 47, 497-509.SUTTON, B. C. & PIROZYNSKI, K. A. (1963)' Notes on British microfungi. I. Trans. Br.

mycol. Soc. 46, 5°5-522.SUTTON, B. C. & PIROZYNSKI, K. A. (1965)' Notes on microfungi. II. Trans. Br. mycol.

Soc. 48, 349-366.VESTEGREN, T. (1900). Eine arktisch-alpine Rhabdospora. Bih. K. suenska Vetensk Akad,

Handl.26, 1-23.

(Accepted for publication 20 February 1967)