Notes on Diplonaevia (Discomycetes inoperculati), with special regard to the species on Juncaceae

Notes on Diplonaevia (Discomycetes inoperculati), with special regard to the species on Juncaceae J. A. Nannfeldt Nannfeldt, J. A. 1984. Notes on Diplonaevia (Discomycetes inoperculati), with spe- cial regard to the species on Juncaceae. - Nord. J . Bot. 4: 791-815. Copenhagen. ISSN 0107-055X. The only monocotyledonous families that take members of Diplonaevia Sacc. (= Merosficfis Clem.) are found to be Juncaceae, Cyperaceae (Caricoideae-Caricieae and Trichophorum) and Poaceae (especially Nardus). The presence or absence of stromatic, clypeus-like tissues is studied in several species and found to be very vari- able not only from species to species but also within individual species. The patterns of variation exhibited by the excipulum, the asci and the spores are also discussed. No less than ten species are known to grow on Juncaceae, eight on Juncus and two on Luzula. D . helicospora (on J . trifidus), D . hyperborea (on Juncus subgen. Alpini) and D . savilei (on L. arcfica) are new species. New combinations are D. anfarcfica (Den- nis) Nannf., D . junciseda (Schroet.) Nannf. and D . microsficfa (Dennis) Nannf. Host ranges and distributions are also studied and discussed. J . A. Nannfeldt, Inst. of Systematic Botany, Univ. of Uppsala, P.O. Box 541, S-751 21 Uppsala, Sweden. Introduction An early attempt (Nannfeldt 1932) to improve the clas- sification of the discomycetes possessing minute, mostly dark, erupment, “mollisioid” apothecia and inhabiting the three “glumiflorous” families (Cyperaceae, Junca- ceae and Poaceae) by combining them into an emended Hysteropezizella v. Hohn. (type species Micropeziza subvelata Rehm) has proved to be of temporary value only, as the resulting genus became too heterogeneous. The same has been the fate of the simultaneously created genus Laetinaevia Nannf. (type species Naevia lapponica Nannf.) correspondingly intended as a “store- house” for species with minute and erumpent, but yel- lowish to pinkish apothecia and mostly inhabiting di- cotyledons. Hysteropezizella became the object of a (far-from- complete) revision by DCfago (1968). She divided it into Hysteropezizella sensu strictiori, Hysterostegiella v. Hohn. (type species Stictis fenestrata Desm. on Scirpus lacustris L.) and Merostictis Clem. (type species Stictis emergens Karst. on Juncus conglomeratus L.) and rele- gated a number of species to Drepanopeziza (Kleb.) v. Hohn., Eupropolella v. Hohn., and Pyrenopeziza Fuck. The splitting of Hysteropezizella sensu meo was decid- edly a step in the right direction, but simultaneously the transfers to Drepanopeziza and Eupropolella resulted in an increased heterogeneity of those genera. Laetinaevia was revised by Hein (1976), who severed Naeviopsis Hein (type species Naevia epilobii Karst.) and relegated a number of species to other genera, e.g. Callorina Korf (= Calloria sensu Tul.) and Ploettnera P. Henn. Hein (1980, 1981, 1982, 1984 and pers. comm.) has later extended his studies to Hysteropezizella and allies, paying special attention to some previously neglected paraphysal characters. By means of them he found it possible to distinguish a “Hysteropezizella-complex” and a “Mollisia- Pyrenopeziza-complex” within the Mol- lisioideae sensu meo, the cardinal mark being the pres- ence of “Paraphysenauflagerungen” in the former and their absence in the latter (cp. Nannfeldt 1984). 0 NORDIC JOURNAL OF BOTANY NORD J BOT 4 791415, MYCOL 046 Nord 1. Bot. 4 ( 6 ) 19R4 79 1 He further restricted Hysterostegiella to the type spe- cies and some likewise small-spored species with lan- ceolate paraphyses thus getting a homogeneous genus, in which Defago’s three large-spored species had n o place. I could then show (Nannfeldt 1984) that H. holo- schoeni (deNot.) DCf. and H. lapponica DCf. (on Tri- chophorum caespitosum (L.) Hartm.) together with some species on cyperaceous, juncaceous and poaceous hosts constitute a new genus, Hysteronaevia Nannf., marked, inter aha, by + Iasci. H. hydrophila (Bomm., Rouss. & Sacc.) Def. (on Scirpus lacustris) belongs to Hysteropezizella according to Hein (1981). The transfer of Naevia paradoxa Rehm (on Juncus trifidus L.) and allies to Eupropolella as performed by Defago has proved to be unjustifiable (Nannfeldt 1983a). They belong instead clearly to the “Hysteropezi- zella-complex”, where they constitute the very natural, resurrected genus Naeviella (Rehm) Clem., marked by I + asci and spores provided with a lateral protuberance topped by a minute germ-pore. Almost from the start I have been strongly impressed by the multitude and diversity of (in part undescribed) minute, k erumpent discomycetes growing on Juncus and - as it seems - to a large extent restricted to single host species, sections or subgenera. Thus the question was asked whether it would be possible to find some correlation between the evolution within the host genus and that amongst its inhabitants, just as is the case with Cyperaceae-Caricoideae and the smuts of the genus An- fhracoidea (cp. Nannfeldt 1979 and the literature cited there, Savile 1979). I t was also realized that the tax- onomic position of less than a third of the named junci- ocolous taxa could be considered as settled. Some of these taxa have been disposed of in two recent papers, viz. one (Nannfeldt 1983a) on the genus Naeviella with, inter alia, N. paradoxa (Rehm) Clem. on J . trifidus, and the second (Nannfeldt 1984) on Hysteronaevia with, in- ter aha, H . clavulifera Nannf. on J . arcticus Willd., J . bi- glumis L . and J . jacquinii L., and H. minutissima (Rehm) Nannf. on J . trifidus and J . gerurdi Lois. as well as H. luzulicola Nannf. on Luzula arcuata (Wg.) Sw. Attention has furthermore been paid to two hitherto mostly neglected features, viz. the individual variation in the location of the apothecia and the changes in the structure of the exipulum during growth, two factors whose importance is illustrated by means of numerous microphotos. The present paper treats the juncicolous and luzulicolous species referable to Diplonaevia Sacc. (= Merostictis). Sundry notes on members growing on other monotyledonous hosts are added. Hein (1984) widens the scope and the host range of the genus by in- cluding a number of species inhabiting dicotyledons, but as they are insufficiently known to me, they will be passed in silence. Unfortunately, the generic name Merostictis (Cle- ments 1909) has to give way to the older Diplonaevia (Saccardo 1889), because Clements (1909) and Cle- ments & Shear (1931; cp. Hein 1976, Sherwood 1977) lectotypified the Saccardoan genus with Stictis caricum Auersw. (? = S . seriata Lib.) and DCfago found this species to be a typical Merostictis, an opinion which Hein endorsed and I now share, though once (Nann- feldt 1932) I considered S. seriata to take a most isolated position, but it and the Nardus-inhabiting M . perpusilla and M. salassorum form a well-marked group within Di- plonaevia because of the shape of their “periphyses”. Clements’ only motive had certainly been the “First Species Rule”, then generally accepted by North Amer- ican botanists, but there is nothing to invalidate his choice. Lambotte (1887) had earlier referred (misunder- stood) S . seriata to his new genus Pseudostictis, but as this genus is now lectotypified (Sherwood 1977) with Peziza punctiformis Pers. (= Stictis pallida Pers.) it be- comes an obligate synonym of Cryptodiscus Corda (1838) and unavailable for S . seriata. In 1932, I suggested that Mollisia luzulina Karst. be- cause of the pinkish colour of fresh apothecia might be- long to Laetinaevia, whereas M. emergens Karst., type of the future Merostictis, then known to me through old herbarium specimens only, was placed in Hysteropezi- zella. In his revision of Karsten’s species of Mollisia, Dennis (1950) followed my tracks, declaring that M. emergens is a good Hysteropezizella and that my sugges- tion about M. luzulina “would seem to be well founded”. DCfago deemed the two species congeneric and consequently transferred M. luzulina t o Merostictis. Defago found the discriminating generic feature of Merostictis to be the texture of the excipulum, its sur- face being smooth, “semblable a celle de certaines Pyre- nomycetes” and formed by a layer of parallel, agglu- tinated, cylindrical and septate cell-rows, all ending at about the same height and with only the terminal cells free from each other. The bulk of the species, in which the excipulum markedly surpasses the hymenium, evi- dently form a most natural entity. Two graminicolous species deviate by apothecia “avec leur marge non dif- fkrenciee et situCe a la hauteur du sommet des asques”. One, M. andropogonis (Berk. & Curt.) DCf., has later been transferred to Nimbomollisia Nannfeldt (1983b) as N. aberrans (Peck) Nannf., and the other M. hypo- pyrrha (Rehm) Def., belongs according to Hein (pers. comm.) to Naeviopsis. Preceded by a nomenclator and a generic description follow here notes on the structure of the apothecia, on the patterns of variation and on the host ranges. These notes confirm the distinctness and the phylogenetical homogeneity of Diplonaevia but offer no clues as to its affinities. A final chapter is devoted to the taxonomy, host ranges and geographical distributions of the species inhabiting juncaceae. 792 Nord. J. Bot. 4 ( 6 ) 1984 Diplonaevia Sacc. Syll. fung. 8: 666 (1x89). - Lectotype (sel. by Clements 1909): Stic/i.s caricum Auersw. (‘? = Stictis sericrm Lib.). Merosriciis Clem.. Gen. fungi p. 64 (1909). -Type: Stictis erner- gens Karst. I’vreuodi.scus Pctr.. Ann. Myc. 25: 202 (1927). -Type: P. cari- cis Pctr. A.stcwnaevicr Petr., Ann. Myc. 27: 40X (1929). - Type A. tri- chophori Pctr. Mostly strictly saprophytic, but a few species ( D . seriata and D. luzulinu) seem to start as parasites but fruit only in dead bleached areas. - Apothecia foliicolous or cul- micolous, scattered to gregarous, immersed, intra- and/ or sub-epidermal, in some (most?) species formed be- neath a “clypeus” of brown-walled cells, which may per- meate the lumina of the epidermis cells and be k firmly connected with the excipulum, minute (0.1Lo.4 mm across), circular t o broadly elliptical in face view, from yellowish to reddish turning to (blackish) red-brown; eventually exposed through cracks in the epidermis, of- ten cross-wise, but not or only little erumpent. at first almost closed with a small apical pore, which gradually enlarges, sometimes through radial slits; excipulum in most species strongly surpassing the hymenium, and the ultrahymenial part when moist dome-shaped to conical, sonictinies even prolonged to a low “chimney”. - Exci- pulum several-layered, outwards of distinct aggluti- nated cell-rows forming a textura angulari-prismatica; the outmost layer reaching the very margin; the ultra- hymenial part inwards producing a kind of “peri- physes”; cell-walls hyaline or those of the external face at least in part brown. - Asci mostly relatively small and slender. cylindrico-clavate, apically rounded, thin- wallcd except for the rather small and narrow I+ apical apparatus. - Spores mostly small, at least in volume, slightly heteropolar, ellipsoid to narrowly cuneate or subacicular, mostly straight, thin-walled, hyaline, often with 2 conspicuous. k subterminal drops best seen in Lactic Blue, in some species eventually with a slightly supramedian septum. only rarely 3-septate. - Para- physes filiform, septate, simple or sparsely branched, hyaline; apical cells slightly enlarged, mostly clavicu- late. rarely k pyriform, only rarely exceeding 2 pm diam. Clypeus-like structures and the location of the apothecia A clypeus as defined by Ainsworth et al. (1971) is a “shield-like stromatic growth, with or without host tis- sue, over one o r more perithecia”, and the last word im- plies that this structure should be restricted t o pyreno- mycetous fungi. I t is true that such structures are most conspicuous and most frequently met with amongst the “Pyrenomycetes”, but they are not totally absent from other groups, e.g. the inoperculate Discomycetes. The presence of a clypeus and its anatomical structure are usually considered as characters on at least generic level. I n the Mollisioidcae scnsu meo superficial apothecia formed beneath a shield of radiating hyphae character- ize the genera Micropeiza Fuck., Nannfeldtia Petrak and Scufomollisiu Nannf. (cp. Nannfeldt 1976). Wolf (1912) described a parasite on roses as the sole member o f a new genus, Diplocarpon, in which the sub- epidermal fruitbodies looked discomycetous, but placed the genus in the Microthyriaceae as the fruitbodies for- med beneath a subcuticular shield of radiating hyphae. Wolf (1924) changed his opinion and referred the genus to the Phacidiales. when he was confronted with a spe- cies without shield but otherwise in essential characters so closely agreeing with the preceding species that he considered them as congeneric. More species were sub- sequently transferred t o Diplocarpon, which is now placed in the Drepanopezizoideae (Nannfeldt 1932, Korf 1973). In one member of Naeviella fungal tissues are found between cuticle and epidermis and also covering the in- ner side of the lid above the apothecium (Nannfeldt Diplonuevia would seem to be a third genus compris- ing both species with and such without clypeus-like structures. Petrak (1927) described as Pyrenodiscus caricis gen. et sp. nov. an ascomycete inhabiting Carex pyrenaica Wg. He found its fruitbodies to possess both disco- mycetous and pyrenomycetous traits, but as they are “oben fast klypeusartig mit der Epidermis verwachsen”, he referred the genus to the Pyrenomycetes, though with hesitation. Two years later, he (Petrak 1929) met with a similar fungus on Trichophorurn caespitosurn and described it as Asteronupvia trichophori gen. et sp. nov. Its fruitbodies were found to have a “mit der Epidermis verwachsene Deckschicht”, which “wohl nur als ein machtig entwickeltes Excipulum aufgefasst werden kann”. He found the genus clearly discomycetous but also very close to Pyrenodiscus, which genus he conse- quently now regarded as discomycetous too. Defago (1968) recognized in them typical members of Meros- ticti.y, i .e. Diplonaevia. Her drawing (DCfago 1968: Fig. 20a) of M. trichophori shows the epidermis ruptured and its cells split but no trace of a “Deckschicht”. A cor- responding drawing (Defago 1968: Fig. 17 left) of M . curicis shows the epidermis ruptured above the centre of the apothecia and the otherwise intact epidermis cells surrounding the rupture permeated with a small-celled fungal tissue; in the text the apothecia are said to de- velop beneath the epidermis “auquel ils sont rattaches par un clypeus”. In M. elynae (Defago 1968: Fig. 21) a thin one-layered clypeus is shown between the intact epidermis and the apothecium but not mentioned in the text. In M . ernergens, M. perpusilla and M. salassorum (Defago 1968: Figs 16. 18, 19) the epidermis is shown as ruptured but otherwise intact, in M . luzulina (DCfago 1968: Fig. 17 right) as ruptured and strongly compressed abovc the apothecium, and in M . circinata (Defago 1983a). SI Nord J R o l 4 ( 6 ) IYX4 793 Fig. 1 . Diplonaevia caricis (Zetterstedt). Excipulum. X ca. 550. - Figs 2-3. D. perpusilla (Holm 533d/71 and J . A . N . 6644a). Ex- cipulum. x ca. 550. - Figs 4-6. D. seriafa ( 4 and 6 Sydow Myc. germ. 408 on Carex hirta; 5 Samuelsson on C. aquarilis x ha- lophila). Apothecia, median sections. X ca. 300,350 and 900, respectively. - Fig. 7 . D. luzulina (J .A.N. 11541). Apothecium, me- dian section. X ca. 350. 794 M o d . J . Bot. 4 (6) 1984 1968: Fig. 20b) the epidermis is partly lost and for the rest split with only the outer wall covering the apothe- cium. Defago thought it possible to distinguish between two groups within the typical members of Merostictis, viz. one ( M . emi~rgens, M . exigua, M . luzulina, M . perpu- silla, M . ,sulassorum, and M . seriuta) with the excipular margin o f textura globulosa and the other ( M . caricis, M . circinuta, M . el,vnae, and M . trichophori) with mar- gin of textura prismatica; an intermediate position was assigned t o M . glucialis: “tres voisine de M . emergens, e l k possi.de une marge dont les cellules polygonales sont quelque peu allongkes, ce qui la rapproche de M . rrichophori et M . circinutu”. I t is easily seen that the drawings referred-to above are schematical and not fully reliable as to the anatomy of the hosts. They furthermore give the impression that, a s ii rule. the host tissues are left almost untouched by weathering and action of the fungus; neither do they show any trace of the mycelium from which the fruit- bodies are born. I t is. indeed. very difficult to get a clear view of the relations of the fruitbodies to the host tis- sues, 21s these are often badly disorganized and as, moreover, extensive comparisons reveal the annoying fact that different fruitbodies. even closely neighbour- ing ones, niay behave differently. Only a few apposite rcsults will be reported here. I t appears that a margin of textura globulosa, as de- scribed by Defago. is not necessarily formed by the fruitbody itself but may be part of a clypeus-like struc- ture adhering to the fruitbody. When the fruitbody ex- pands. this covering tissue cannot follow and bursts eventually into rfr triangular flaps exposing the proper margin o f the underlying fruitbody or. if the covering is firmly attached to the excipulum, they may burst to- gether causing the margin to become lacerate. ( I ) In D. caricis (Fig. 1) the excipular cells are thus relatively short but distinctly prismatic and mostly hid- den beneath a covering o f dark-walled. f isodiametric. angular cells. ( 2 ) D. perpusilla (Figs 2-3) agrees on the whole but a dark-celled covering is only occasionally found. (3) D. emergens s l a t . (i.e. incl. M . glacialis and Tro- chila juncicolu) with its much wider host-range has been studied more in detail because of its great variability, a variability that at least in part is influenced by specific differences in the anatomy of the culms. The epidermis, ruptured and lifted by the developing apothecia may remain intact. but its cells are often com- pressed or split and deprived of the basal wall, which may disappear or remain joined with the subepidermal tissues. The cell-lumina may be permeated and almost filled by hyphae, and this in unbroken cells as well as in such without the basal wall. These hyphae correspond evidently to the “clypeus” in D. caricis and may adhere to the apical part of the apothecium. On J. conglomeratus (the type host) and its close akin J. effusus the fruitbodies are mostly flat-bottomed, though only exceptionally so flat as that drawn by De- fago (1968: Fig. 16), which rests upon unchanged meso- phyll. But even in those rare cases, when the mesophyll cells have kept their shape, the cells and intercellulars of at least two adjoining layers are permeated by hyphae, not shown in her drawing. In fact. the apothecial base is mostly bulging and compresses the underlying cells (Figs 8-9). sometimes the base may even be broadly conical or hemispherical penetrating into the substrate and pushing its cells aside. O n the other hand, apothecia with hemispherical base are the rule on J. filiforniis (Fig. l o ) , and this led DCfago to refer the fungus on this host to M . glacialis (type host J . trifidus). The variable shape of the apothe- cia1 base on the two first hosts combined with their close affinity to J. filiformis makes this distinction fallacious. The largest apothecia (up to 0.4 mm diam) are to be found on the two first more robust hosts, whereas those o n the tiny J. filiforrnis hardly exceed 0.15 mm. They arc also apt to remain small on thc more occasional hosts. The larger apothecia may remain open even when dry with the yellowish hymenium still flat and the flaps o f the epidermis and the ultrahymenial excipulum rccurved; the pigmentation of the excipular cell walls is usually faint, but yet distinct. Smaller and darker apo- thecia niay occur on the same culms as large ones, but thcn as ii rule in separate areas, evidently produced by myixlia yielding o n l y small fruitbodies. Could this be caused by later infections. when the culms have already lost most o f its nutrients? I n addition to the apothecial shape three more marks are considcrcd to discriminate M . glacialis from D . ernergens, viz. the dark colour of the upper part of the opothecium, the rarity o f septate spores and the struc- ture o f the margin. The type collection of M . glacialis (Figs 11-14) shows apothecia with a rounded base and a thick, strongly pigmented “clypeus” hiding the margin of the cxcipulum, which lower down is partly concealed by ;I close wcb of winding hyphae, but the differences are not sharp enough for regarding these fungi as speci- fically distinct. (4) In D. luzulina (Fig. 7) the apothecia may be cov- ered by compressed epidermal cells, but the covering mostly consists of the outer halves only of split such cells, and, in both entire and split cells, hyaline- to brown-walled hyphae as a rule permeate the lumina. ( 5 ) D. seriata is either a very variable polyphagous species or, perhaps rather, an aggregate of a few allied species (cp. Defago 1968: 49-50), but the two samples treated here seem to be typical of the species sensu stric- tissimo. In the sample on Carex hirta (Figs 4, 6) the apothecial covering is mostly formed by entire epider- mal cells, which are slightly compressed and permeated by brown-walled hyphae, but sometimes the basal wall is lost, and occasionally parts of sclerenchyma strands are incorporated into the cover. In that on C. aquatilis X halophila (Fig. 5 ) the covering epidermal cells are empty and may be slightly compressed. It remains an 51’ Nord. J. Bot 4 ( 6 ) 19R4 795 open question whether the difference is due to the fun- gus or to the host. (6) Sections of D . circinata on Juncus “articulatus” (Fig. 17) show erumpent apothecia loosely covered by remnants of the disintegrating epidermis, and a col- lection on J . triglumis (Fig. 18) has apothecia almost completely covered by the strongly disintegrated epi- dermis. In neither case are covering hyphae observed. (7) D . junciseda (Figs 25-29) deviates from all the preceding species by the shape of its apothecia, which are not depressed but broadly ampulliform, penetrate deep into the palisade tissue, pushing its cells aside and, in a relatively small area only, touch the epidermis, which is ruptured and may in part be thrown off. The apothecia are upwards and often laterally covered by a ? coherent crust of thick- and dark-walled fungal tissue. The excipulum does not or only little surpass the hyme- nium. (8) In D . exigua (Figs 32-37) the apothecia are of about the same shape as those of the preceding species but the ultrahymenial part may be extremely long. The early stages are well described by Desmazieres (1847): “La Sticris exigua annonce d’abord sa presence par une fente, le plus souvent ovale ou allongee, entouree, comme une petite collerette, de I’epiderme blanche. Au fond, sont nichees de tres petites cupules, qui ne parais- sait bien que par suite de I’humectation prolongee du support. Ovales ri I’etat sec, elles s’ouvrent et s’arron- dissent de plus en plus par I’humidite, mais e lks restent toujours enfoncees et concaves. Leur disque est d’une fauve Claire et ses bords sont noiritres.” After maturing the apothecia disintegrate rapidly leaving the pale- walled cylindrical holes open. No hyphae can be seen in the epidermal cells surrounding the ruptures. The struc- ture of the excipulum is less regular than in any of the species treated above; its individual cells are rather an- gular than prismatic but nevertheless form distinct rows. The excipulum proper seems at least in part to have a covering of dark-walled fungal tissue. The excipulum proper The apothecia of all species studied possess at the base an external excipulum of textura globuloso-angularis, and this changes laterally into a textura angulari-pris- matica or, more rarely, a strict textura prismatica. The zone where this change takes place varies a little from species to species, and in the upper (peri- and ultra-hy- menial) part there are notable interspecific differences as to size (length and breadth) and shape of the cells as well as in thickness and pigmentation of the cell walls, differences that mostly are difficult or impossible to ex- press intelligibly in words but that, with some experi- ence, make several of the species unmistakable under the miscroscope. Most peculiar is D . trichophori (Figs 38-39), where the radiating, relatively narrow cell-rows are formed by cells with straight longitudinal walls and scanty septa at exactly right angles. In the great major- ity of species the cells are a little angular or barrel- shaped, which makes the cell-rows distinctly constricted at the septa and bulging between them, as shown by the photos of D . caricis (Fig. l ) , D . perpusilla (Figs 2-3), D . emergens (Figs 13-14), D . circinata (Figs 19-21, 40-44), D . savilei (Figs 22-24), D. juncisedu (Fig. 29), D . exigua (Fig. 37), D . hyperborea (Figs 4 5 4 8 ) , and D . micro- stictcl (Figs 53-54). The external layer may be so transparent that the layer next below becomes visible. Its cell-rows are as a rule more slender, the cells less bulging and the walls paler. In some species the cells forming the margin initially possess thick and strongly refractive (“glassy”) walls, and in D . savilei (Figs 22-23), D . circinata (Figs 40-44) and D . hyperborea (Figs 46-48), where the margin long remains in this stage, the refractive walls are very con- spicuous. The ultrahymenial part of the exipulum is on the in- side clad with free periphysis-like hyphal ends. These structures are delicate, and it is mostly difficult or even impossible to study them in detail on herbarium mate- rial, otherwise they would often afford useful specific characters. Suffice to note that in D . litzulina (Defago 1968: Fig. 17 right) these free ends reach a length of 18- 30 ym and may be septate, whereas in D . perpusillu (Defago 1968:Fig. 18). D . salassorum (Defago 1968: Fig. 19) and D . seriata (Hein 1984) they may be almost isodiametric and only ca. 4 ym long. In the taxonomic part, the descriptions of the exci- pulum will be kept rather short and are admittedly not comparable to each other. Asci In comparison with related and similar genera the asci (and the spores) of Diplonaevia are on the average small. The largest asci are those of D . trichophori (Fig. 50) , which may reach 75 x 14 pm. The asci are on the whole rather uniform and offer little of interest. The apical apparatus is small and I + , though in some in- stances the reaction is faint or even vacillating. The asci are often slender, but rather plump in e.g. D . caricis, D . elynae and D . trichophori; the stipe is as a rule short and indistinct. They often taper conically into a k truncate tip (e.g. in D . elynae, D . exigua and D . seriata), result- ing in so characteristic an arrangement of the spores that Minks (1882: 201) a century ago found it worthy of recording: “. . . die Anordnung der Sporen eine nach der Figs 8-12. Diplonaevia emergens (8 and 9 type of N . obscure-marginara on Juncus effusus, 10 Holm 330 on J . filiformis, 11 and 12 type of P. glacialis on J . rrifidus). Apothecia, median sections. x ca. 700, 900, 550, 550 and 700, respectively. 796 Nord I Bot 4 (6) 198.1 Figs 13-16. Diplonaevia emergens. -Figs 13-14 (type of P. glacialis). Excipulum. X ca. 600. -Figs 15-16 (type of T. juricicolu on J . gerardi). Excipulum and hymenium. X ca. 900. Figs 17-21. Diplonaeviu circinufa. - Fig. 17 (type). Apothecium, median section. X ca. 350. -Figs 18-21 (J .A.N. 811 on J . biglu- mis). Apothecium. median section, X ca. 300 , and excipulum. X ca. 900. -Figs 22-24. D. savilei. -Figs 22-23 (Wickhom). Mar- gin of excipulum. x ca. 600. - Fig. 24 (Haglund & Kallstrom). Excipulum. X ca. 900. 798 Nord J Bot 4 ( 6 ) 1984 Figs 32-37. Diplonaevia exigua (Figs 32-35 type, Figs 3 6 3 7 type of Naevia rehrnii). - Figs 32-36. Apothecia, median section, note the abnormal, double-sided “Janus”-apothecium in Fig. 35. Figs 32 and 34 X ca. 350, Figs 33 and 35-36 x 550. - Fig. 37. Exipu- lum, x ca. 550. Figs 25-31. Diplonaevia junciseda. - Figs 25-28 (Holm 337). Apothecia, longitudinal sections. X ca. 350, Fig. 28 x ca. 600. - Fig. 29 (Torne Lappmark Asplund). Excipulum. X ca. 600. - Figs 3&31 (Andersson, PI. lapp. 222) . Spores. x ca. 900. Nard I Bot 4 (6 ) 1YR4 80 1 Basis hin zusammengedrangte, wahrend in die Spitze meist nur eine Spore zu liegen kommt”. The number of spores is normally eight but tends to become reduced. In some species ( D . antarctica, D . caricis and D. trichophori) four is the only or dominant number, in others (e.g. D. circinata and D. savilei) 4- spored asci are k frequent. Spores As mentioned above the spores of Diplonaevia are on the whole comparatively small or, more exactly, of com- paratively small volume, for there are species in which the spores are very long but then at the same time very narrow, subacicular. They are in principle heteropolar but often almost imperceptibly so. Part of the species have narrowly calviculate to sub- acicular spores with the contents homogeneous or, a t the most, enclosing a few very small droplets and either permanently 0-septate or eventually I-3-septate. These species can according to increasing spore-lengths be ar- ranged as follows: D. perpusilla (spores 5-7 pm long); D. luzulina ((& )1&13(-16) pm); D. emergens ((10-)12-17(-24) pm, Figs 15-16); D. salassorum (12-18 pm); D. caricis (17- 28 pm); and D . helicospora (55-70 pm, Figs 51-52). The last-mentioned species exposes a most excep- tional feature; before reaching maturity the spores grow much too long for the asci and roll up into a densely coiled fascicle, but on ejaculation they unroll and turn straight or slightly curved or sigmoid. The mature spo- res are consistently 3-septate with almost equally long segments. The spores of D. emergens and D. luzulina may even- tually get a faint. slightly supramedian septum, but this seems to happen only occasionally and in single apothe- cia, in which case, however, the number of septate spo- res may be rather large. In other species the spores are likewise slender and the contents homogeneous or at the most enclosing a few droplets but narrowly fusiform and straight to sub- falcate in shape. Ranked according to increasing spore- length they are: D. hyperborea (14-20(-25) pm, Fig. 49); D . micro- slicta (20-29 pm, Figs 53,SS-56); and D . junciseda ((28- )3(k35(-42) mm, Figs 3C31). In D . junciseda the mature spores get a slightly su- pramedian septum, but in the two other species they re- main 0-septate. A third group of species is marked by much plumper spores, 2.5-6 pm wide, oblong-ellipsoid to subcylindric with rounded ends, the contents with 2 or more large drops and often a number of droplets, and only excep- tionally getting a faint septum. The following species belong here: D. seriata (6-10 x 3 pm); D. circinata (8-12 x 3 pm); D. savilei (9-15 X 3 pm); D. antarctica ((10-)12-16 x 2.5-3 pm, Fig. 57); and D. trichophori (18-26 x 4-6 pm, Fig. SO). As a rule the intraspecific variation of the spores keeps within rather narrow and well-defined limits as to both size and shape, which makes their diagnostic value high. Only in D . exigua (Fig. 36) is the shape more irreg- ular and “untidy”, but this may serve as a character too. The spores (size 15-20 x 3-6 pm) d o not fit any of the groups distinguished above. They are mostly oblong-cu- neate and tending to become fusiform, the contents are mostly spumose, large, occasionally confluent drops are sometimes k distinctly seen. Paraphyses The paraphyses of Diplonaevia are on the whole rather uniform and trivial, offering little of interest. I t may, however, be observed that in D . hyperborea the tips have a tendency to become more strongly and more abruptly enlarged than usual. Only D. exigua deviates more markedly. Besides be- ing unusually numerous its paraphyses have very nar- row lumina and thick “glassy” walls, are, as it seems, upwards very slightly widened and conglutinated in a way that obstructs detailed observations. Patches are of- ten found, in which the hymenium is “powdered” by oval cells (about 3 X 2 pm or slightly larger) and where similar bodies are to be found in the hymenium attached to hyphae. Rehm (1888: 163), who described the para- physes as apically rounded and up to 5 pm across, evi- dently observed these bodies too, but I cannot decide whether the paraphyses may act as conidiophores or a parasitic hyphomycete has infected the hymenium. Some remarks on host ranges and distributions It is obvious that only part (perhaps only a minor part) of the existing species of Diplonaevia and similar genera connected with juncaceous hosts are known at yet. Such species seem to occur all over the world, though most frequently at high altitudes and high latitudes, and to a large extent to possess narrow host ranges. Further- more, it should not be forgotten that our present knowl- edge is highly influenced by the circumstance that, in the phanerogamic herbaria, specimens of some Juncus- species regularly comprise also overwintered parts lia- ble to house these fungi, whereas specimens of other Figs 38-39. Diplonaevia rrichophori (38 Berta Anderson 279b. 39 type). Excipulum. x ca. 350 and 550, respectively. - Figs 4&44. D. circinata (Figs 40 and 42 J .A.N. 24024 on J. biglumis, Fig. 41 Disko, Fries on J. biglumis and Figs 4344 Hedberg 3128 on J. triglumis). Excipulum, Figs 40 and 42 the same spot with changed focus. Figs 43 and 44 ditto. All x ca. 550. 802 Ncird J Bot 4 (6) 19x4 species (e.g. J . conglomeratus, J . effusus and J . filifor- mis) are not accompanied by such remnants. The best known areas are Fennoscandia and the Arc- tic islands, but even these are insufficiently known as manifested by single odd samples of unknown species. Two species with comparatively wide host ranges are D . luzulina on Luzula subgenera Luzula, Anthelaea and Pterodes and D . emergens on Juncus subgenera Genuini, Septati, Alpini, and Pseudotenageia. Their Eu- ropean distributions are also wide, even if they are markedly rarer in the lowlands and it is doubtful whether they reach the mediterranean zone. A single find of D . luzulina in Siberia shows that its distribution extends far east. Within their host ranges certain hosts are obviously more susceptible than others, thus L . pi- losa is the preferred host of D . luzulina, and some mem- bers of Juncus subgen. Genuini, viz. J . conglorneratus, J . effusus, J . filiformis, and (?) J . jacquinii, are prefer- red by D . emergens. D. circinata is another species with a wide host range on Juncus, but not coincident with that of D . emergens, as subgen. Genuini is excluded, and members of Septati and Alpini are preferred hosts. There are two species, whose host ranges are re- stricted to single subgenera of Juncus, viz. D . hyper- borea to subgen. Alpini, i.e. J . biglumis, J . castaneus, and J . triglumis, and D . exigua to some (but not all) members of subgen. Septati or, at least, J . anceps and J . subnodulosus. There are finally species growing on single host spe- cies, viz. D . savilei on L. arctica, D . junciseda on J . arc- ticus (incl. J . a. x filiformis), and J . helicospora on J . trifidus. It should be observed here that the distribution of D . junciseda is restricted (northern) in comparison with that of its host, being common in the North but so far unknown from central and south Sweden, Denmark, Britain and continental Europe. D . helicospora, unmis- takable because of its most peculiar spores, is so far known only from a few adjacent localities in north- ernmost Sweden. Its host is generally placed in subgen. Pseudotenaneia but takes a very isolated position, de- viating “especially in the structure of its leaves. The ser- rulate leaf margin is quite unique for the genus and is reminiscent of related genera as Prionium and Luzula. The lacerate auricles and the occurrence of lacerate bracteoles are also features unusual in the genus Jun- cus.” (Nilsson & Snogerup 1972). I t should not pass unnoticed that J . trifidus in addi- tion is the exclusive or principal substrate for some al- lied fungi, viz. Hysteronaevia minutissima (Rehm) Nannf. (also on J . gerardi; Nannfeldt 1984), Naeviella paradoxa (Rehm) Clem. (Nannfeldt 1983a), and two species of doubtful affinities, Naevia junci Rehm (= Hysteropezizella j . Nannf.) and Mollisia subconica Rehm (= Pyrenopeziza s. Sacc.). The total host range of Diplonaevia on monocotyle- dons is highly thought-provoking, restricted as it is to the three “glumiflorous” families and its occurrence on each of them offering special, important and interesting problems. The four species on dicotyledons that Hein (1984) transfers to Diplonaevia, on the other hand, at- tack four widely scattered host-families. It can no longer be doubted that Cyperaceae and Jun- caceae are closely related, whereas there is a wide gap between them and Poaceae; yet they are all considered to belong t o the same superorder, Commeliniflorae (cp. e.g. Dahlgren & Clifford 1982). Savile (1979; 464) finds their parasitic fungi to support these views and gives Claviceps as an example of a genus of parasites re- stricted to them. For obvious reasons saprophytes are on the whole less strictly substrate-bound, but taxa lim- ited to the same three families are e.g. Hysteropezizella diminuens, the genus Hysteronaevia (Nannfeldt 1984) and the present genus. (1) Though future studies on a global scale will in- crease the number of species of Diplonaevia, there is no reason to believe that Juncaceae will loose its position as the principal host-family. This combined with the fact that its species possess precise and individually different host ranges makes the conclusion almost inevitable that the connection between Diplonaevia and Juncaceae was established very long ago and that since then speciation has worked parallel in them, but this does of course not mean that the birth-place of Diplonaevia must have been on a member of Juncaceae. There is, as far as I am aware, only one more genus relative to which Juncaceae plays an important, though not so dominating role, viz. Clathrospora Rbh. s.str. (Wehmeyer 1961). a genus whose total host range, but for some accidental “jumps”, is the same “glumiflo- rous” families. (2) In Cyperaceae the species of Diplonaevia are re- stricted to Caricoideae-Cariceae (Carex, Kobresia) and Trichophorum of Scirpoideae-Scirpeae, i.e. exactly the same constellation as is the total range of a genus of bio- trophic parasites, Anthracoidea (Kukkonen & Timonen 1979, Savile 1979: 486487). The species on Trichophorum, A . scirpi (Kuhn.) Kukk., is morphologically primitive and takes a rather isolated position. This together with rust relationships and the fact that Trichophorum, alone among the Scir- peae, possesses a Carex-type embryo has led to the sug- gestion “that Caricoideae sprang from very close to Tri- chophorum” (Savile 1979). This suggestion may gain some additional support by D . trichophori, which takes an isolated, though not necessarily primitive position Figs 4 5 4 9 . Diplonaevia hyperborea. -Figs 45-48 (Fig. 45 Lynge, Fig. 46 Rosenvinge, Fig. 47 Kjellman & Lundstrom, and Fig. 48 Wulff). Excipulum. All x ca. 550. - Fig. 49 (Wiren). Asci and spores. X ca. 900. - Fig. 50. D. trichophori (Berta Anderson 279b). Hymenium. X ca. 900. 804 Nord. I. Bat. 4 (6 ) 1984 within its genus, being extreme in two characters, viz. the structure of the excipulum and the dimensions of asci and spores. (3) In Poaceae Nardus strictu L. hosts two species of Diplonaevia, the common and widespread D. perpusilla (Rehm) Hein (= Trochila exigua Rostr.) and the rare D. salassorum (Def.) Hein, known from the type col- lection only. N. stricta is a most distinctive plant, generally treated ;IS forming a monotypical tribe (Nardeae). It is obvious that i t does not belong to the festucoids, and it seems to be so different from all other grasses by a number o f re- markable and fundamental features that i t ought to be given a much higher (subfamily’?) rank. The seemingly distichous spicate inflorescence is in fact tristichous: all spikelets but one are arranged alternately in two of the rows and the uppermost spikelet alone represents the third row. The pistil possesscs a single style continuing in ;I filiform papillate stigma. The fruit is trigonous. Narrlus is far from attractive to fungi, even such of groups that otherwise are richly represented on grasses; i t takes n o smut, no rust, no mildew, either downy or powdery, no Phyllachora and no Septoria or septorioid fungus. In thc last moment for being inserted here Dr. Hein informed me that, on some other grasses, fungi occur that he considered as belonging to the D. seriatm com- plex and gave me the opportunity to study them, but I havc not been ablc to arrive at a definite decision about their taxonomic position. Key to the species of Diplonaevia on Juncaceae I . On L u z i r i r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 I . On Juncus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Spores slendcr. (&)I& guttate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. D. luzulina 2 . Spores oblong-claviculate 3(15) x 3(-3.5) pm, 4 3. Spores longer, slender.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 4. Spores narrowly claviculate, (l(k)l2-17(-24) X 1.5-2(-2.5) pm. eguttate. rarely 1-septate . . . . . . . . . . . 3. D. emergens 4. Spores plumper, 2 . 5 4 pm wide. mostly with 2 large drops, 5. Spores 8-12 x 3-4 pm . . . . . . . . 6. Spores (I(k)12-16 x 2.5-3 pm. . . . . . . . . . . . 1. D. untarctica spored . . . . . . . . . . . . . . . . . 7 7. Apothecia depressed, exposed through expanding pores in the epidermis. Spores 14-20-25) x 2 . 5 4 pm. On J . biglu- mis, J . castaneus and J . triglumis . . 7. Apothecia ampulliform, sunken in d longitudinal mouth. Spores 15-20 x 3 4 pm. On J. anceps and J . subnodulosus.. . . . . . . . . . . . . . . . . . . . . 4. D. exigua 8. Spores 20-29 x 3 pm. narrowly fusiform, 0-septate. (South Georgia). . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. D. 8. Spores more than 30 pm long. 1-3-septate .. . . guttate. On L. urctica . . . . . . . . . . . . . 10. D. savilei 3. Spores up to 20 pm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . 0-scptate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5 . Spores longer . . . . . . . . . . . . . . . . . . . . . . . . . . 6 9. Spores (28-)3&35(42) X 2.5-3 pm, I-septate, not coiled. On J . arcticus and J. a. X filiformis. . . . . . 7. D. junciseda 9. Spores 55-70 X 1.5 pm, 3-septate, coiled. On J . trifidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . D. helicospora 1. Diplonaevia antarctica (Dennis) Nannf. comb. nov. Hysteropezizella antarctica Dennis, Kew Bull. 22: 445 (lY68). - Type: on Juncus sp.; South Georgia F.I.D.S. (= Falkland Is- lands Dependencies Survey) 3048 (K!). Maw.: Juncus sp. Exsiccata: None. Illustrations: Fig. 57. Apothecia culmicolous, scattered to gregarious, ca. 0.2- 0.25 mm diam. - Excipulum basally of textura globu- losa; peri- and ultra-hymenially of textura angulari-pris- matica of very delicate, 3-4 ym broad cellrows; cells with thin, only faintly pigmented walls. - Asci saccate to subcylindrical. 40-50 x 5-7 ym, truncate, I + , (almost always) only 4-spored. - Spores (mostly obliquely) l-se- riate, oblong, (10-)12-16 X 2.5-3 ym, straight, with 2 large drops and often 2 very small subterminal droplets, 0-septate. - Paraphyses filiform, septate, simple, with pyriform heads, ca. 2 pm diam. The spores of D. antarctica agree rather well with those of D. circinata but are markedly longer, and the exci- pulum does not show the “glassy”-walled margin so con- spicuous in D. circinata. Occirrrrnce: Known so far only from South Georgia (2 col- lections). 2. Diplonaevia circinata (Lib.) Hein Sydowia 36: 101 (1984). Stictis circinata Lih.. PI. crypt. Ard. n . 232 (1834). - Nuevia cir- cirratu Rehm in Rbh., Krypt.-FI., ed. 2, 1.3: 144 (1888). -Me- rosrictis circinata Def., Sydowia 21: 54 (1968). - Type: on Jun- cus crcutiflorus; Belgium, Lib. crypt. Ard. n. 232. Stictis lamyi Mont. in sched. - Cryptodiscus lamyi Rbh., Bot. Zeit. 9: 454 (1851; nom. nud.). - Naevin lamyi Rehm in Rbh., Krypt.-FI., ed. 2, 1:3: 145 (1888). - Hysreropezizella lamyi Nannf.. N . Acta Reg. Soc. Sci. Upsaliensis4:8:2: 118 (lY32). - Spec. auth.: on J. articulatus; France H“-Vienne Limoges Lamy (K!, S-Rehm!. S-Syd.!). Mollisiu alpina Rostr.. Meddr Gronland 3: 609 (1891). -Type: on J. alpinus; Greenland Igaliko Rosenvinge ( C ! ) . Matr.: Juncus acutiflorus Ehrh.. J. alpinus Vill. s.lat., J. al- pinus X articulatus, J . ariiculatus L . , J . biglumis L., J . castan- eus Sm., J . gerardi Lois., and J . triglumis L. s l a t . (i.e. incl. J. albescens (Lge) Fern.). Exsiccara: Lib., PI. crypt. Ard. 232 ( J . acutiflorus; “S. cir- cinata”, type). -PI. vasc. groenl. exs. 40 ( J . castaneus, on dead culms together with D. hyperborea). Illustrations: Figs 17-21,40-44. - Defago 1968: Figs 2e- f , 151, 20b. Figs 51-52. Diplonaevia helicospora (type). Hymenium. X ca. 900. - Figs 53-56. D. microsticfa (Figs 53-55 type, Fig. 56 Greene 1496). Figs 53-54. Excipulum. x ca. 550. Fig. 55. Hymenium. X ca. 900. Fig. 56. Spores. X ca. 1800. - Fig. 57. D. anfarctica (Greene 1632). Hymenium. X ca. 900. 806 N o d . J . Bot. 4 ( 6 ) 1984 Apothecia culmicolous or foliicolous, scattered to gre- garious (in sufficiently stout culms up to ab. 50 apo- thecia crowded into rounded spots, a few mm across, or into narrow girdles encircling the culms), subepidermal, 0.2-0.3 mm diam, (blackish) brown, at maturity ex- posed through longitudinal or cruciate slits in the epi- dermis, only little erumpent when moist and retracting on drying. - Excipulum peri- and ultra-hymenially few- layered of textura angulari-prismatica; the external layer with thick brown cell-walls, the underlying layers with rather thick “glassy” walls. - Asci cylindrico-clavi- culate. 35-50 x 5-10 pm, apically rounded, I + , often 4- spored. - Spores 1-seriate, when 8 2-seriate or irreg- ularly arranged, cylindrico-oblong, 8-12 X 3-4 pm, with 2 large subterminal drops, 0-septate. - Paraphyses fili- form, septate, simple or moderately branched, apically claviculate, up to 2 pm across. According to Defago (1968: 55) a conidial state of Ce- phnlosporium-type is formed in cultures as well as thick- walled “chlamydospores”. Microscopically, D. circinatu is easily known by the short, cylindrico-oblong. 2-guttate spores. An unmis- takable feature visible (when present) to the naked eye is the circinate apothecial groupings, which gave the species its epithet. This is well shown by most samples on members of Juncus subgen. Septati. On the other hand, the morphologically exactly agreeing apothecia on members of subgen. Alpini hardly group that way, and so the possibility was seriously considered that they might be specifically distinct. But this idea had to be abandoned, when it gradually turned out that, even in Septati, distinctly crowded apohecia are to be found on culms of a certain stoutness only and that they are evenly dispersed on more slender organs such as inflo- rescence branches. Nevertheless, the host-subgenera will be kept apart in the following listing of the material known to me. Occurrence: D . circinata is widely distributed being known from the Central European lowlands as well as from the Alps, Fennoscandia. Arctic Eurasia and Arctic North America. It is not common except on J . biglumis and J . triglumis and is, like most of its congeners, decidedly favoured by high altitudes and high latitudes. The finds on Juncus subgen. Septati are from the following areas: -on 1. acutif7oru.T (?) from Belgium (type of S. circinatu) and Germany (Westfalen, Defago 1968, Hein 1984). -on J . al- pinus from Sweden (Jamtland Frason, UPS!), Fennoscandian USSR (Lapponia ponojensis, UPS!). Norway (Hedmark Fol- dal, UPS!; Oppland Jotunheimen. UME!; and Sor-Trondelag Kongsvoll. UPS!), France (H“-Savoie, Defago 1968). Swit- zerland (K‘ Graubiinden and K’ Wallis, Defago 1968). Italy (Val d’Aosta. DCfago 1968). Greenland (South- and West- greenland. type of M. alpina and 4 more finds, Rostrup 1891, C!. UPS!). Newfoundland (Straits of Belle Isle, UPS!), -on J . alpinus X articulatus from Switzerland (K’ Graubiinden Ber- gun Miiller, UPS!, ZT!), -on J . articulatus from Sweden (Upp- land, UPS!), Norway (Hedmark Tynset. UPS!), France (HIc- Savoie. Difago 1968; and H“-Vienne, type of S. lamyi, K!, S!, UPS!. Rehm (1888) gives “Genfer-See” (i.e. Lat. “Lacus Le- manus”) as the locality having evidently misinterpreted the “Limovicum” (i.e., Limoges) of the label), Switzerland (K’ Graubunden, Defago 1968; UPS!), and Austria (Steiermark, GZU!, UPS!). The finds on Juncus subgen. AIpini are as follows: -on J . bi- glumis from Novaya Zemlya (UPS!). Svalbard (Vestspitsber- gen, Lind 192th as B. juncisedum, O!), Sweden (Hiirjedalen, Jamtland, Pite Lappmark and Torne Lappmark, UPS!), Nor- way (Hordaland, Oppland and Finnmark, UPS!), Greenland (Westgreenland 3 finds, UPS!), Canada (Distr. of Franklin 3 finds, DAOM!, UPS!), and U.S.A. (Alaska, UPS!). - on J . cas/aneus from Greenland (Westgreenland Holsteinborg Distr.. C!, UPS!, and Eastgreenland Charcot’s Land, PI. vasc. groenl. exs., UPS!), and Canada (Distr. of Keewatin South- ampton Isl.. DAOM!), - on J . triglumis from Svalbard (Vest- spitsbergen 5 finds, Lind 1928a as Naevia pusilla, O! UPS!), Sweden (Harjedalen. Norrbotten, Lule Lappmark and Torne Lappmark 15 finds in all, GZU!, UPS!), Norway (SGr-Tronde- lag and Finnmark, UPS!), Finland (Lapponia enontekiensis. UPS!). Switzerland (K’ Graubunden, UPS!), Siberia (2 finds. UPS!), and Greenland (West- and Eastgreenland 3 finds, A single find on J . (subgen. Pseudotenageia) gerarcii is from UPS!). Sweden (Norrbotten Ncder-TorneB, UPS!). 3. Diplonaevia emergens (Karst.) Hein Sydowia 36: 84 (1984). Stictis emergens Karst., F. fenn. n. 765 (1868); Not. Sallsk. F. FI. Fenn. Forh. 11: 254 (1870). - Mollisia emergens Karst., My- col. fenn. 1: 194 (1871). - Trocbila emergens Rehm, Hedwigia 22: 35 (1883). - Naevia emergens Karst.. Acta Soc. F. FI. Fenn. 2:6: 168 (1885). - Pbragmonaevia emergens Rehm in Rbh.. Krypt.-FI., ed. 2, 1 3 : I64 (1888). - Pyrenopeziza emergens Boud.. Discom. d’Eur. p. 133 (1907). - Merostictis emergens Clem., Gen. fungi p. 64 (1909). - Hysteropezizella emergens Nannf.. N. Acta Reg. Soc. Sci. Upsaliensis 4:8:2: 115 (1932). - Type: on Juncus conglomeratus; Finland Karsten (F. fenn. n. P.seudopeziza glacialis Rehm, Ascom. n . 818 (nom. nud.); Hedwigia 24: 232 (1885). - Pbragmonaevia glacialis Rehm in Rbh., Krypt.-FI.. ed. 2, 1:3: 164 (1888). - Diplonaeviaglacialis Sacc., Syll. fung. 8: 667 (1889). - Hysleropezizella glacialis Nannf.. N. Acta Reg. Soc. Sci. Upsaliensis 4:8:2: 1 I6 (1932). - Merostictis glacialis Def., Sydowia 21: 45 (1968). -Type: on J . trifidus; Italy Trentino-Alto Adige Rehm (Ascom. n. 818). Trocbilajuncicolu Rostr., Bot. Tidsskr. 15: 231 (1886). -Type: on J . “compressus” (= gerardi); Norway Troms KBfjord Warm- ing (C!). Stictis niesslii Roumeg.. Rev. myc. 9: 24 (1887). - Type: on J . effusus; (lectotype here selected) Czechoslovakia Niessl (Rou- nieg., F. gall. 3936a). Naevin obscure-marginata Starb., Bih. K. Sv. Vet.-Akad. Handl. 21:3:5: 18 (1895). -Type: on J . effusus; Norway Akers- hus Driibak StarbPck (S! holotype, UPS! isotype). Matr.: Juncus alpinus Vill., J . arcticus Willd. s l a t . (i.e. incl. J . balticus Willd.), J . arcticus X filiformis, J . articulatus L.. J . ca- stanei~s Sm.. J . conglomeratus L., J . effusus L., J . filiformis L., J . gerardi Lois., J . jacquinii L., J . squarrosus L., and J . trifidus L. s la t . (Le. incl. J . bostii Tausch). Exsiccata: Jaap, F. sel. 106b (1. effusus; “Naevia pusilla”). - Karst., F. fenn. 765 ( J . conglomeratus; “S. emergens” type). - Krypt. exs. vindob. 2028 ( J . conglomeratus; “Pbr. emergens”). - Lund. & Nannf.. F. exs. suec. ined. ( J . filiformis). - Petr., FI. boh.-mor. 1:l: 1871 ( J . sp.; “Pbr. emergens”). - Rbh., F. eur. 2829 ( J . effirsus; “S. exigua”). - Rehm, Ascom. 657 ( J . effusus; “T. emergens”), 818 ( J . trifidus; “Ps. glacialis” type). - Rou- meg., F. gall. 3936a ( J . effusus; “S. niesslii” lectotype), 3936b ( J . effusus; “S. niesslii” paratype). - Syd., Myc. germ. 2153 ( J . effusus; “Phr. emergens”), 2154 ( J . conglomeratus; “Pbr. emergens”), 2539 ( J . effusus; “Pbr. emergens”). - Non: Petr., Mycoth. gen. 201 1 (Carex sempervirens; “Phr. emergens”), q.e. Diplonaevia sp. nov. (?). 765). Nord J Rot 4 (6) 19x4 808 Illustrations: Figs 8-16. - Rehm 1888: 123, Figs C 1-5. - Starbiick 1895: Figs 4a-b. - Defago 1968: Figs Id , 15a, d , e. 16. Apothecia culmicolous or foliicolous, subepidermal, 0.1-0.4 mm diam, with flat to subhemispherical or con- ical base, yellowish to dark brown, exposed through mostly cruciate slits, not or only little erumpent, re- tracting on drying. - Excipulum many-layered, peri- and ultra-hymenially of textura angulari-prismatica, the cells of all layers shorter (almost isodiametric) towards the margin. - Asci cylindrico-claviculate, 35-50 X 5-7 pm, I + , 8-spored. -Spores 2-seriate, narrowly cuneate, (1&)12-17(-24) x 1.5-2(-2.5) pm, straight or faintly curved, eguttate, when fully mature often with a slightly supramedian septum. - Paraphyses filiform, septate, apically claviculate, up to 2 pm across. The crucial mark of D. emergens is the slender, nar- rowly claviculate, eguttate spore, sometimes getting one, slightly supramedian septum. The apothecium is mostly rather pale with shades of yellowish to ochrace- The scope of the species is here widened to include Pseudopeziza glacialis, Trochila juncicola and Naevia obscure-marginata, whereby also its host range becomes much wider. As mentioned above, the shape of the apothecia obviously depends upon the anatomy of the host; thus they are as a rule rather flat-bottomed on J . conglorneratus and J . eflusus, whereas their base is hemispherical to conical on the related J . filiformis. The latter shape also characterizes the mostly darker apo- thecia on J . trifidus, which host takes a very isolated po- sition taxonomically. Fungi with exactly agreeing spores and on the whole matching morphology but for various amounts of pig- mentation also occur on J . alpinus, J. arcticus, J . articu- latus, J . castaneus, J. gerardi, J . jucquinii, and J . squar- rosus. The taxonomic value of Trochila juncicola, described on J. gerardi from northernmost Norway, long re- mained doubtful t o me. as no supplementary material on this host could be found in spite of eager search in the UPS phanerogamic collections and in the field. The situation changed last autumn, when Mrs and Dr Holm brought me four collections from the very type area and it could be established that they in all respects match D. emergens. The name Trochila juncicola was, in several later pa- pers, loosely applied by Rostrup to various species of Hysteropezizella s. lat. on Juncus and Luzula. My revi- sion of available samples gave the following results, Rostrup 1888: Naeviella paradoxa on J . rrifidus; Ros- trup 1891 : Hysteropezizella pusilla on J. arcticus (Iga- liko); Blytt & Rostrup 1891: H. diminuens on J . arcticus and J . trifidus, and D. savilei on L . arctica; Rostrup 1903: D. emergens on J. filiforrnk; and Rostrup 1904: D. luzulina on L. pilosa. ous. Lind (1910a. b, 1924b, 1926, 1928a. b, 1934), a com- patriot of Rostrup’s and his successor as the authority on Arctic micromycetes, took Rostrup’s species in the same wide and loose sense but found it necessary to change the name into the older Naevia pusilla (Lib.) Rehm, as he considered also this to fall within his com- posite “species”. Occurrence: D. ernergens has a pronouncedly montane and northern (but not arctic) distribution. I t cannot be decided whether it shows some physiological specialisation, but it is evi- dent that some hosts arc preferred; thus J . arcticus, J . con- glorneruius, J . effusus, J . filiformis, and J . jacquinii, all be- longing to subgen. Genuini, are frequently infected. The same may apply t o J . trifidus in the Alps. but there are no finds of it in the north. The limitation of the finds on J . gerardi to the pro- vince of Troms in north Norway is noteworthy and so far inex- plicable. The finds on J . alpinus, J . ariiculaius and J . squar- ro.sus are scarce and scattered. In northern Europe it is so far known from Sweden (Upp- land on J . arciicus UME! and on J . filiforrnis UPS!; Harjedalen o n J . filiforrnis UPS!; Jamtland Are on J . articulaius J.A.N. 11656d UPS! and on J . casianeus K. Johansson UPS!; Vast- crbotten on J . filiforrnis Lund. & Nannf. F. exs. suec.; Nor- rbotten on J . urcticus x filiforrnis UPS!; Asele Lappmark and Pitc Lappmark on J . arcticus UPS!; Lule Lappmark on J . arcri- (‘us UPS! and Jokkmokk on J . alpinus J.A.N. 12044b UPS!; Tbrne Lappmark on J . arcticus GZU!, UPS!), from Finland (Satakunta on J . arciicus X filiforrnis UPS!; Tavastia australis on J . conglornermtus, type of S. ernergens, and on J . filiformis H!; Ostrobottnia borealis on J . arciicus, Defago 1968; Lappo- nia kemensis on J . urciicus x filiforrnis UPS!; Lapponia en- ontekiensis on J . Jliforrnis UPS!), from Fennoscandian U.S.S.R. (Lapponia ponojensis on J . arcticus H!), from Nor- way (Akershus on J . effusus, type of N . obscure-rnarginata; Nordland on J . arcticus x filiforrnis UPS!; Troms on J . arcticus UPS!. o n J . filiforrnis UPS! and on J . gerardi, type of T. junci- cola, 5 finds C!, UPS!) and from Iceland (on J . arciicus UPS! and on J. filiforrnis, Rostrup 1903 as T. juncicola, Larsen 1932 as N . pusilla, C!). The Central European area as sketched below is based on finds from Germany (Thuringen on J . effusus, Jaap F. sel., S!, and Oberhof on J . squarrosus, Jaap 429 as Phr. glacialis S!; Sachsen Erzgebirgeon J . effusus S!; Bayern Bayerischer Wald on J . efjcusus and J . conglorneraius, Syd. Myc. germ. 2153 and 2154). France (H“-Garonne on J . effusus, Roumeg. F. gall. exs. 3936b; HIc-Savoie on J . effusus, Jaap 1907, S! , and on J . jucquinii UPS!; Alpes-Maritimes on J . irifidus UPS!, ZT!), Switzerland (K’ Graubiinden on J . arciicus GZU!, UPS!, ZT!, o n J . filiforrnis UPS!, ZT!, on J . jacquinii UPS!, ZT!, on J. iri- fidus, Defago 1968 as M. glacialis, GZU!, UPS!, and Rha- tische Alpen Dischmatal Teufli, on J . ariiculatus Scheuer GZU!, UPS!; K‘ Wallis on J . jacquinii UPS! and on J . irifidus, Defago 1968 and Miiller 1977 as M. glacialis), Austria (Tirol on J . jacyuinii UPS!; Karnten on J . effusus GZU!, UPS! and on J . irifidus, Rehm 1912 as Phr. glacialis, S!; Steiermark on J . effu- sus GZU!, UPS!; Nieder-Osterreich on J . conglorneraius, Krypt. exs. Vindob., S!) , Italy (Piemonte on J . irifidus, Defago 1968 as M. glaciulis; Trentino-Alto Adige on J . trifidus, Rehm Ascom. 818. type of P. glacialis, Rehm 1888, 1912), Czechoslo- vakia (Morava on J . conglorneratus S! and on J . effusus, lecto- type of S. niesslii, Petr. FI. boh.-rnor., Rbh. F. eur., Rehm As- com. 657, Roumeg. F. gall. exs. 3936a). and Poland (Slask on J . effusus, Syd. Myc. germ. 2539). The fungus should be sought for in Britain, Asia and North America. 52 Nord J . H o t 4 ( 6 ) 1984 809 4. Diplonaevia exigua (Desm.) Hein Sydowia 36: 93 (1984). Stictis exigua Desm., PI. crypt. Fr. I: 1533 (1847); Ann. Sci. Nat. Bot. 3%: 183 (1847). - Phragmonaevia exigua Rehm in Rbh., Krypt.-FI., ed. 2, 1:3: 163 (1888). - Hysteropezizella exi- gua Nannf., N. Acta Reg. SOC. Sci. Upsaliensis 4:8:2: 115 (1932). - Merostictis exigua Def., Sydowia 21: 46 (1968). - Type: on Juncus “articulatus” (? = anceps); France Calvados Roberge (Desm., PI. crypt. Fr. I: 1533). Naevia rehmii Jaap, Verh. Bot. Ver. Brandenburg 47: 83 (1905). - Hysteropezizella rehmii Nannf., N. Acta Reg. SOC. Sci. Upsaliensis 4:8:2: 120 (1932). - Type: on J . anceps var. atricapillus; Denmark SGjnderjylland Jaap (F. sel. 83). Matr.: Juncus anceps La Harpe s.lat. (i.e. incl. J . atricapillus Drejer), (?) J . articulatus L., and J . subnodulosus Schrank. Exsiccata: Desm., PI. crypt. Fr. I: 1533 (1. “articulatus”; type of S. exigua); 11: 1033 ( J . “articulatus”; “S. exigua”). - Jaap, F. sel. 83 (J. anceps; type of N. rehmii). - Moug. & Nestl., St. crypt. vog.-rhen. 1241 ( J . “articulatus”; “S. exigua”). - Rou- meg., F. gall. 338 ( J . “articulatus”; “S. exigua”). - Westend., Herb. crypt. 939 (n.v.). - Non: Rbh., F. eur. 2829 (J. effusus; “S. exigua”), q.e. D . emergens. - Syd., Myc. germ. 944 ( J . con- glomeratus; “Phr. exigua”), q.e. Hysteropezizella pusilla (Lib.) Nannf. Illustrations: Figs. 32-37. - DCfago 1968: Fig. 1% - Dennis 1960: P1. xx, Fig. I; 1968a: PI. xxiii, Fig. I; 1978: PI. xxvii, Fig. I . Apothecia culmicolous, ampulliform, deeply immersed, 0.2-0.4 mm across, externally dark (reddish-blackish) brown, with hymenium pale fulvous, exposed through short longitudinal slits, which gradually widen into ellip- tic - circular pores, not or only little erumpent. - Exci- pulum several-layered, peri- and ultra-hymenially of textura angulari-prismatica, the cell-walls of the out- most layer brown, darkest towards the margin. - Asci cylindrico-claviculate, 60-80 x 8-12 pm, mostly conica- lly tapering into a rather narrow k truncate tip, I + , with relatively thick, highly refractive walls, 8-spored. - Spo- res apically as a rule 1-seriate and for the rest 2-seriate, rather variable in shape, oblong-cuneate to fusoid or rather irregular, 15-20 X 3-6 pm, with spumose con- tents and often large, sometimes confluent drops, (ac- cording to Dennis) eventually 1-septate. - Paraphyses numerous, filiform, inconspicuously septate, with thick, light-refractive walls, moderately branched, upwards slightly claviculate, sometimes pyriform, up to 3 pm across. D . exigua is at first sight known from all other micromy- cetes on Juncus by the short slits in the culms, which lead into deep cylindrical holes enclosing the apothecia. Microscopically it is marked by the size of the spores and, especially, by the abundance of paraphyses and their thick, refractive walls. It obviously takes a most isolated position within the genus, but no better posi- tion can be suggested. Occurrence: D . exigua is inconspicuous and evidently much overlooked. Its so-far-known area is very restricted, west-Eu- ropean, but it seems to be fairly common regionally. The east- 810 ernmost find is that on the Baltic island of Gotland. There may be some doubt about its host range. The only proven hosts are Juncus anceps and J . subnodulosus, and it is most unlikely that it grows on J . articulatus in its recent restricted sense. The finds known to me are the following: Sweden: Gotland, Lummelunda Kinners ( J . subnodulosus) 27 Jul 1973 Thedenius (UPS!). - Denmark: S~jnderjylland, R B ~ B Isl. Lakolk ( J . an- ceps) 16 Jul 1904 Jaap 253 (S!), 18 Jul 1903 Jaap (F. sel. exs., type of N. rehmii). - Britain ( J . subnodulosus, Dennis 1960, 1968a, 1978). - Belgium: West-Vlanderen, between Ostende and Ravenszyde (J. “articulatus”) July 1847 Desmazieres 989 (DCfago 1968). - France: sine loco ( J . “articulatus”) hyeme Mougeot & Nestler (St. crypt. vog.-rhen.). Dtp‘ Calvados, Caen ( J . “articulatw”) Roberge (UPS-Fries!). Hermanville ( J . “articulatus”) July Roberge (Hb. Barb.-Boiss. 1074; S!); Au- gust. - (Desm., PI. crypt. Fr., type of S. exigua). Lyon-sur-Mer ( J . “articulatus”) Roberge (S!, UPS!). - Germany: Bayern, Oberbayern, Kr. Starnberg S. of Pocking ( J . subnodulosus) 30 Jul 1967 Poelt 4918a & 11501 (GZU!, UPS!). 5. Diplonaevia heliocospora Nannf. sp. nov. Apothecia initio subepidermalia, ca. 0.1-0.15 mm diam, Asci subcylindrici, 50-65 x 5-7 pm. Sporae fas- ciculatae, fasciculis ad postremum spiraliter contortis, subaciculares, 55-70 X 1.5 pm, 3-septatae. Type: on Juncus trifidus; Sweden Tome Lappmark Holm 1070a (UPS!). Matr.: Juncus trifidus L. Exsiccata: None. Illustrations: Figs 51-52. Apothecia hypophyllous or culmicolous, scattered, 0.1- 0.15 mm diam., hymenium eventually exposed through a small rupture in the epidermis and a brown-walled “clypeus”, on drying completely retracting, blackish to- bacco-brown. - Excipulum several-layered, peri- and ultrahymenially of textura angulari-prismatica, outmost layer of relatively narrow (3-3.5 pm) rows of only slightly bulging cells, margin k hidden by rather dark- walled, angular and * isodiametric “clypeus” cells. - Asci slender, subcylindrical, 50-65 x 5-7 pm, I + , 8- spored. - Spores fascicled, before maturity growing considerably longer than the asci causing the fascicles to coil strongly but uncoiling on ejaculation, subacicular, 55-70 X 1.5 pm, with tapering ends and 3 equidistant septa. - Paraphyses relatively sparse, filiform, septate, apically very narrowly claviculate, up to 2 pm across. D. helicospora is easily known by its long, filiform, 3- septate spores, before ejaculation forming strongly coiled fascicles. Occurrence: This outstanding and easily observed species must be rare. It is known from four finds only, but they are rather rich and emanate from an area less than 10 km across, viz. Sweden: Tome Lappmark Kiruna (formerly Jukkasjarvi) Kerkevagge, reg, alp. 18 Jul 1952 L. Holm 1069 (UPS!); ditto nr. Lake Ripsajaure, reg. alp. 18 Jul 1952 L. Holm 1070a (type). Katterjaure Railway Halt 26 Jul 1952 L. Holm 1120 (UPS!). Between Riksgransen Railway Station and the River Katterjokk 20 Jul 1953 L. Holm 1202 (UPS!). N o d J . Bot. 4 (6) 1984 6. Diplonaevia hyperborea Nannf. sp. nov. Apothecia subepidermalia, erumpentia, ca. 0.1-0.2 mm diam., initio melleo-flava, deinde obscure brunnea. Sporae fusiformes, 14-20(-25) x 2 . 5 4 pm, guttulatae, 0-septatae. Paraphyses filiformes, clava saepe pyriformi ad 3 vm diam. Type: on Juncus biglumis; Greenland John Murray Isl. Wulff (UPS! holotype, C! isotype). Matr.: Juncus biglumis L., J . castaneus Sm, and J . rriglumis L. Exsiccata: PI. vasc. groenl. exs. 40 ( J . castaneus, on dead eulms together with D. circinata). Illustrations: Figs 4 5 4 9 Apothecia follicolous or culmicolous, scattered, subepi- dermal, 0 . 1 4 . 2 mm diam, erumpent through longitudi- nal slits in the epidermis, initially honey-yellow, turning dark reddish brown. - Excipulum several-layered of rather thin- and pale-walled cells; basally of textura glo- buloso-angularis; peri- and ultra-hymenially of textura angulari-prismatica of cell-rows up to 7 pm broad and septate into ca. 8-12 pm long cells; marginal cells ini- tially truncate with thick ”glassy” walls. - Asci cylin- drico-claviculate to saccate, 4MO X 6-12 pm, apically obtuse, (often indistinctly) I+ , 8-spored. - Spores irreg- ularly clustered to obliquely 2-seriate, fusiform, 14-20 (-25) x 2 . 5 4 pm, straight, inequilateral to faintly curved, mostly guttulate, 0-septate. - Paraphyses fil- iform, simple or occasionally branched, septate, to- wards the tips claviculate or more abruptly widened to a pyriform head, up to 3 pm diam. Characteristics of D. hyperborea are the pale and large- celled excipulum and the relatively large, fusiform spo- res. Occurrence: D. hyperborea is confined to three species of Jun- cus subgen. Alpini, viz. J . biglumis, J . castaneus, and J . triglu- mis. On J . biglumis it is so far known only from the Arctic coast of Siberia and from the islands north of Europe and America, polewards reaching 82”45’N (John Murray I d . ) . On J . castaneus it has been found also in the mountains of U.S.A. (Colorado) and China (Sikang). On J . triglumis there is only one find (Greenland). The samples studied by me are the following: Novaya Zem- lya: Lichutin Isl. (= “Barents 0”) ( J . biglumis) 16 Aug 1921 Lynge (Lind 1924b as Naevia pusilla, O!). Fretum Matochkin ( J . biglumis) 7-13 Jul 1875 Kjellman & Lundstrom (UPS!). - Svalbard: Vestspitsbergen, Treurenbergfjorden Point Crozier ( J . biglumis) 21 Jul 1899 Wulff (UPS!); Longeyeardalen ( J . castaneus) 6 Aug 1924 Lid (Lind 1928a as Belonidium juncise- dum. O!); Isfjorden Bohemans tundra ( J . biglumis) 6 1 4 Aug 1921 Wiren (UPS!); Sorkapplandet ( J . biglumis) 19 Aug 1920 Lid (Lind 1928a as N . pusilla, O!), - Greenland: Northgreen- land, John Murray Isl. ( J . biglumis) 3 Jul 1917 Wulff (Lind 1924a as B. juncisedum type; C!, UPS!; Kap York, Insygigsak ( J . biglumis) 26 Jul 1883 Nathorst (UPS!); Murchinson Sound, Igdloluarsuit ( J . biglumis) 20 Aug 1921 Nygaard 11: 66 Fung. 2 ( C ! as R. juncisedum det. Lind, UPS!. The record in Lind 1926 of a find with the same date but from “Neke” may be due to some confusion, as no voucher can be found, and may refer to the present collection). - Eastgreenland, Hold-with-Hope ( J . biglumis) 20 Jul 1891 Hartz (UPS!); Jamesons Land ( J . casta- neus) 5 Aug 1891 Hartz (UPS!) ; Charcots Land (71”54’N, 29”W) 450 m s.m. ( J . castaneus) 6 Aug 19.58 Einarsson & Hol- men (PI. vasc. groenl. exs., UPS!); Scoresbysund, Sydkap-Oen (71”20’N, 25”S’W) ( J . biglumis) 13 Aug 1927 Stwensen 273 Fung. (UPS!). - Westgreenland, Disko “in locis sterilibus uligi- nosis ad latera inferiora alpium” ( J . biglumis) Jul 1833 Vahl (UPS!), Lyngmarken ( J . castaneus) 4 Jul 1871 Fries (UPS!), ditto 25 Jul 1886 Rosenvinge (C! as Mollisia junciseda det. Ro- strup); Christianshaab ( J . castaneus) 26 Jul 1884 Warming & Holm (Rostrup 1888 as M. junciseda; C!, UPS); Itiflak ( J . tri- glumis) 7 Aug 1883 Berlin (UPS!). - Southgreenland, Sydest- bugten, inland of the peninsula Anguertussoq ( J . biglumis) 14 Aug 1932 Grentved 253 Fung. (UPS!). - Canada: Distr. of Franklin, Ellesmere Isl. Fram Harbour ( J . biglumis) 4 Aug 1899 Simmons 1400 Fung. (UPS!); Baffin Isl. Frobisher Bay ( J . castaneus) 1 1 Aug 1959 Hedberg 3167 Fung. (UPS!). - U.S.A.: Colorado, Clear Creek Co. Mt. Evans 13500’s.m. ( J . castaneus) 23 Aug 1959 Weber & Willard 10923 Fung. (UPS!), 15 Aug 1960 Weber, Porsild & Holmen (UPS!). - U.S.S.R.: Arctic Siberia, Dickson Harbour (73”28’N, 80”58’E) ( J . biglu- m u ) 6 1 0 Aug 1878 Kjellman (UPS!); Konyarn Bay ( J . casta- neus) 2&30 Jul 1879 Kjellman (UPS!). -China: Sikang, Taofu (= Dawo) Distr. between Ngata and Takuan ea. 3900 m s.m. ( J . castaneus) 11 Sep 1934 Smith 12049 Fung. (UPS!). 7. Diplonaevia junciseda (Schroet.) Nannf. comb. nov. Trochila junciseda [“(Karst.?)”] Schroet., Jahresb. Sehlesis- chen Ges. Vaterl. Cultur 58: 169 (1881); Hedwigia 21: 151 (1882). - Hysteropezizella junciseda Nannf. in Lund. & Nannf., F, exs. suec. Fasc. 47-48: 48 (1956). -Type: on Juncus arcticus; Sweden, Lule Lappmark Lake Pollaure 6 Aug 1856 Wichura (n.v.). Topotype: on dead culms in Anderss., PI. lapp. 222. Matr.: Juncus arcticus Willd. s.lat. (i.e. incl. J . balticus Willd.) and J . arcticus x filiformis L. Exsiccata: Anderss., PI. lapp. 222 ( J . arcticus, on dead culrns). - Lund. & Nannf., F. exs. suec. 2394 ( J . arcticus; “ H . junciseda“) . Illustrations: Figs 25-31. Apothecia culrnicolous, immersed, broadly ampulli- form, 0.241.3 mm across, long covered by the epider- mis, then exposed by radiating slits, not or hardly erum- pent, blackish brown. - Excipulum not or only little sur- passing the hymenium, perihymenially of textura angulari-prismatica, outmost layer of cells 5-10 x 3 4 pm, in a marginal zone of ca. 75-100 pm breadth with strongly pigmented cell walls but lower down much paler, marginal part often to some extent covered by a k coherent crust of dark cells. - Asci plump, cylindrico- claviculate, (50-)5540(-80) x 9-1 1(-14) pm, roundtip- ped, I + , 8-spored. - Spores fasciculate, subacicular to subfusiform, straight, slightly curved or subsigmoid, slightly heteropolar with the upper half only little thicker than the lower, (2%) 30-35(42) X 2.5-3 pm, rather early with a distinct median septum, exception- ally 3-septate, at maturity sometimes breaking into the two halves. - Paraphyses simple, filiform, septate, only slightly enlarged upwards, reaching ca. 1.5 pm diam. D. junciseda stands out from all its congeners by its spo- 811 52’ Nord. J . Bot. 4 ( 6 ) 1984 res: slender, mostly 30-35 pm long, straight to sub- falcate, early with a very distinct median septum, strongly resembling those of Hysteronaevia advena (Karst.) Nannf. This species was established in a rather bewildering way. In his account of the fungi collected by Wichura in North Sweden Schroeter (1881: 169) lists a Mollisia jun- ciseda Karst. on leaves of Eriophorum russeolum, but his short description gives full evidence that this fungus was instead Mollisia advena Karst. (= Hysteronaevia advena). This entry is immediately followed by : “Tro- chila juncisedu (Karst.?) - Auf Halmen von Juncus arc- ricus Willd. - Gesellig iiber einen grossen Teil des Hal- mes verbreitet. Becher etwa 0 . 2 4 . 3 mm breit, ganz ein- gesenkt, beim Anfeuchten nicht hervortreten, schwarz- lich. Schluche fast cylindrisch, 5@55: 10 p, 8-sporig. Sporen stabchenformig, 33-35: 2.5-3 p. oben oft etwas breiter. Inhalt in der Mitte meist mit einer Scheide- wand. Paraphysen fadenformig, 1 breit. - In seinen mikroskopischen Verhdtnissen der vorigen fast gleich. Vielleicht ist es nur von der Beschaffenheit der Nahr- pflanze bedingt, o b die Becher aus der Oberhaut vortre- ten oder von ihr bedeckt bleiben, es wiirde dann ein so erheblicher Unterschied einzelner Mollisia-Arten (Nip- rera), die zu den Pezizen, und Trochila-Arten (Naevia), die zu den Stictideen gerechnet werden, nicht bestehen. - Am See Pollaure, 6. August 1856. - Vielleicht identisch mit der von Karsten in Finnland gefundenen Mollisia juncinella *junciseda K.”. Schroeter had here changed the epithets, for in fact Karsten (1869, 1871) placed juncinella as a subspecies under junciseda (not vice versa). Some years later, when listing a fungus on Eriopho- rum from northernmost Norway as M . junciseda-jun- cinella Schroeter (1888: 273) gave the epithets in the correct sequence but a few pages later (p. 280) he ex- hibits another proof of absentmindedness by reporting M. junciseda on Greenlandish “Juncus caespitosus”, for thre is no such Juncus, it was probably Scirpus caespito- sus that he had in view. Schroeter’s arrangement of Wichura’s two fungi and his concluding remark are unintelligible, unless we as- sume that his intention was to connect the one with jun- ciriella and the other with junciseda (but which with which?). Be this as it were, 7‘. junciseda must be re- garded as a validly published name for the fungus on J . arcticus. Although no authentic material has been avail- able, there can be no doubt about its application, for the detailed description exactly matches a fungus common on this host in the type area. As t o Karsten’s taxa Dennis (1950) has shown that M. junciseda should be placed in Belonopsis (Sacc.) Rehm and that *juncinella “superficially and structu- rally” closely resembles the former. If we accept his la- ter interpretation of the genus Nipteru Fr. (Dennis 1972), they clearly belong to that genus and are, in my opinion, only doubtfully distinct from the younger N . pulla (Phill. & Keith) Boud. Occurrence: D . junciseda has a pronouncedly northern distri- bution. Almost all samples seen by me emanate from the phanerogamic collection in UPS. Its distribution in the Scandes (mainly on ssp. arcticus) agrees fairly well with that of this sub- species (Hulttn 1971: map 437), reaching southwards to Opp- land in Norway and to Harjedalen in Sweden, but from the coasts, where it grows on ssp. balticus (Hulttn 1971: map 440). it is known in the north only. The Swedish finds are north of 64”20’N (Vasterbotten Lovinger, Norrbotten Pitea and Hapa- randa). It is not known from Finland, and the only Norwegian coast locality is Nordland Andoya (on J . arcticus X filiformis). In Iceland it seems to be common (on ssp. intermedius Hyl.). It has further been seen from Westgreenland (Sukkertop- pen, Rostrup 1888: 538 as M. junciseda, C!, and Godthaab) as well as from north Canada (Manitoba Fort Churchill and Distr. of Keewatin Southampton I d . ) . There are so far no finds either from Britain or from Conti- nental Europe. 8. Diplonaevia luzulina (Karst.) Hein Sydowia 36: 86 (1984). Mollisia luzulina Karst., Myc. fenn. 1: 194 (1871). - Naevia lu- zulinu Karst., Acta Soc. F. F1. Fenn. 2:6: 168 (1885). - Pha- cidium luzulinum Lamb., FI. myc. Belgique Suppl. 1: 265 (1887). - Phragmoriaevia luzulina Rehm in Rbh., Krypt.-FI., ed. 2, 1:3: 163 (1888). - Laetinaevia luzulina “Nannf.” in Lind, Kgl. Danske Vid. Selsk., Biol. Medd. 11:2: 93 (1934). - Mero- stictis luzulina Dtf. , Sydowia 21: 47 (1968). -Type: on Luzula pilosa; Finland Ostrobottnia australis Vasa 22 Jul 1867 Karsten (Hb. Karsten 4007. H. holotype; S-Hb. Rehm!, S-Hb. Sydow!, UPS! isotypes). Naevia luzulae Sacc., Bull. Soc. Bot. Ital. 1893: 18 (1893). - Type: on L . pilosa; Siberia Mt. Borus Martianoff 2254 (PAD!). Matr.: Luzulu alpinopilosa (Chaix) Breistr. (= L . spadicea (All.) DC.), L . arcuata (Wg.) Srn. s.lat. (i.e. incl. L . confusa Lindeb.), L. luzuloides (Lam.) Dandy & Wilm. (= L. albida (Hoffm.) DC. = L. nemorosa (Poll.) E. Mey.), L. multiflora (Retz.) Lej., L. nivea (L.) DC., L. pallescens Sw. x sudetica (Willd.) DC. (= L . “multiflora X sudetica” sensu Lindb. fil.), L . parviflora (Ehrh.) Desv., L. pilosa (L.) Willd., L . silvatica (Huds.) Gaud. (= L . sieberiTausch), L. sudeticu (Willd.) DC., L . wahlenbergii Rupr., and possibly also L. spicafa (L.) DC. Exsiccata: Krieg., E sax. 736. - Petrak, Myc. gen. 154. - Rehm, Ascom. 1010. - Syd., Myc. germ. 2369. - All on L. pi- losa as Phr. luzulina. Illustrations: Fig. 7. - Naumov 1964: Fig. 46. - D6fago 1968: Figs ISf, 17 right-hand half. Apothecia scattered, subepidermal or intraepidermal, flat-bottomed, mostly epiphyllous in dead bleached ba- sal parts of over-wintered leaves, sharply set off from the green, still living distal parts, rarely culmicolous, 0 . 2 4 . 4 mm across, reddish and mostly surrounded by a dark brown rim of dead host cells permeated by hyphae with pigmented walls, exposed through slits in the epi- dermis mostly in the shape of four triangular lobes, re- tracting on drying. - Excipulum peri- and ultra-hyme- nially of textura angulari-prismatica, cells with rela- tively pale walls, the periphysis-like hyphae unusually long, 18-30 pm, and often septate. - Asci cylindrico- claviculate, obtusely pointed, I + , 35-50 x 4-6 pm, 8- 812 Nord. I. Bot. 4 (6 ) 1984 spored. - Spores obliquely 2-seriate, narrowly cuneate, (6-)10-13(-16) x 1.5-2 pm, eguttate, sometimes even- tually with a slightly supramedian septum. - Paraphyses filiforrn. septate, apically not or only faintly enlarged, 1.5-2 pm across. D . luzulina is certainly close to D . emergens but differs by growing on Luzula, by the reddish colour of the fruitbodies and by the unusually long, septate peri- physis-like hyphae on the inside of the ultrahymenial excipulum. Occurence: D. Iuzulina is widely distributed and attacks sev- eral broad- and soft-leaved Luzulae (subgenera Pterodes and Anthelaea). especially L . pilosa. Morphologically indistin- guishable and probably conspecific fungi occur on some narrow-leaved Luzulm too, though more rarely. The species, which seems to start its growth as a parasite. is best developed in latc spring and early summer. It is decidedly more frequent towards the north and in the mountains. In the Nordic countries i t is known on L.pilosa from almost thc wholc of Sweden (Smiland, Ostergotland, Sodermanland, Nlrke. Uppland, Vistnianland, Halsingland, Angermanland, mark and Tome Lappmark, Nannfeldt 1928, Lind 1934, S!, UPS!) but is evidently rare in the southern provinces. It is fur- ther known from throughout Finland (Nyland. Satakunta, Ta- vastia australis, Ostrobottnia australis, Kuusamo, Lapponia keniensis. and Lapponia inarensis; Karsten 1871, type of M. luzulina. H!. S!, UPS!) and from Norway (Akershus and Nord- land, Rostrup 1904 as Trochilm juncicola, Nannfeldt 1928, O!, UPS!), but so far neither from Denmark nor from Iceland. In north Sweden i t has now and then been collected on L. muhiflorrr (Hlrjedalen and Jamtland. UPS!) and on L. par- viflora and 1,. wahlenhergii (Torne Lappmark, Nannfeldt 1928. Lind 1934. UPS!); there is also a record on L. spicata (Torne Lappmark, Lind 1934). In Finland it has been found on L. ar- c u m (Lapponia inarensis, UPS!), L . multiflorn (Savonia au- stralis, H!), and L. pallescens x suderica (Savonia borealis, H!) as well as in Fennoscandian USSR on L . arcuata (Lapponia tu- lomensis, UPS!) and L . parviflora (Lapponia murmanica, H!), There is also a record of it from Novaya Zernlya on L. ar- cuara (Lind 1034). I t is not known from the British Isles but is found on L. pi- losa in various parts of continental Europe: Belgium (Lam- botte 1887), Germany (Westfalen, Petr. Myc. gen., Syd. Myc. germ.; Bayern. Hein 1984; and Sachsen, Krieg. F. sax., Rehrn Ascom.), Poland (Slask, Schroeter 1893) and USSR (Lenin- grad obl., Naumov 1964 as N. luzulae and f h r . luzulina) and further in Siberia (Mt. Borus (ca. 53”N, 87”E). Saccardo 1893, type of N. luzulae). In the Centraleuropean countries it has been found on the broadleaved L. alpinopilosa in Switzerland (K‘ Wallis, Defago 1968, Miiller 1977), L. luzuloides in Austria (Nieder-Oster- reich, S!; and Steiermark. GZU!), L. nivea in Italy (“Siid-Ti- rol”, Rehm 1912; S!) , L. silvafica in Germany (Bayern, Hein 1984) and in Switzerland (K’ Wallis, Dtfago 1968, Miiller 1977), as well as on the narrow-leaved L . sudetica in Swit- zerland (K’ Wallis, Miiller 1977). There is a report on L. parviflora from Greenland that seems reliable. viz. one by Lind (1934). but an older one by Rostrup (1888), “Sp. 1&20 x 2-3 I(” is very dubious. Other- wise it is not known from the Western hemisphere. H:’ q e d a l e n . ’ JBmtland, Vasterbotten, Norrbotten, Pite Lapp- 9. Diplonaevia microsticta (Dennis) Nannf. comb. nov. Hysteropezizella microsfirfa Dennis, Kew Bull. 22: 447 (168). - Type: on Juncus sp.; South Georgia, F.I.D.S. 1504 (K!) . Marr.: Juncus sp. Exsiccara: None. Illustrations: Figs 53-56. Apothecia culmicolous, scattered to gregarious, 0.1- 0.16 mm diam. - Excipulum basally of textura globulosa of cells ca. 3 4 pm diam, and with strong brown walls; peri- and ultra-hymenially of textura prismatica of 3-5 pm broad cellrows, septate into 5-8 pm long, brown- walled cells. - Asci cylindrico-saccate to clavate, 40-50 x 7-9 pm, roundtipped, I + , 8-spored. - Spores clus- tered, narrowly fusiform, 2G29 x 3 ym, straight, inequilateral or faintly curved, slightly heteropolar, eguttate, 0-septate. - Paraphyses filiform, septate, sim- ple, apically slightly claviculate, up to 2 pm diam. The spores (size and shape) are sufficient to discrimi- nate D . microsticta from all its congeners. Occurrence: Known so far only from South Georgia (2 col- lections). 10. Diplonaevia savilei Nannf. sp. nov. Hysreropezizella pusilla sensu Savile in Grovs & Elliott 1971. Diplonaevia circinatu peraffinis sed differt sporis graci- lioribus, paullo longioribus et distincte clavulatis, 9- 13(-15) x 3(-3.5) pm, et habitatione in Luzula arctica. Type: on Luzula arctica; Sweden, Torne Lappmark 2 Jul 1939 Smith (UPS). Matr.: Luzula arctica A. BI. (syn. L. nivalis sensu Beurl. et auctt .). Exsiccata: None Illustrations: Figs 22-24. Apothecia mostly epiphyllous, scattered, subepidermal, ca. 0.1-0.25 mm diam., erumpent through small, gradu- ally widening holes, at first pale brown, then turning blackish brown. - Excipulum peri- and ultra-hymenially of textura angulari-prismatica, the external layer of cells ca. 4-5 pm broad, up to 10 pm long and with relatively thick, brown walls; the marginal cells with thick “glassy” walls. - Asci saccate to subcylindrical, 40-55 (- 70) x 6-9 ym, apically tapering into the obtuse or al- most truncate tip, I + , mostly 8-spored. - Spores mostly obliquely 2-seriate, oblong-claviculate, (9-)10-13(-15) x 3(-3.5) pm, guttate, mostly with one large drop to- wards each end, sometimes with several smaller and ir- regularly arranged drops, 0-septate. - Paraphyses fili- form, septate, mostly simple, upwards claviculate and sometimes slightly torulose, up to 2.5 pm diam. Nord. 1. Bot 4 (6) 1984 813 D . savilei is easily known from D . luzulina, the other Diplonaevia on Luzula, by the structure of the excipular margin and by the plumper and guttate spores. Occurrence: D . savilei seems to attack L . arctica only but to be rather common on it and to follow it everywhere. These are the finds known to me: Novaya Zemlya: mainland E. of Lichutin Isl. 17 Aug 1921 Lynge (Lind 1924b as N . pu- silla, O!). - Svalbard: Vestspitsbergen, Isfjorden Green Bay s. of the Aldegonda Glacier 20 Jul 1915 Asplund (UPS!); Bell- sund 1924 Lid 33 (O! as H . pusilla, det. Lind). - Norway: Troms, Kifjord Mt. Birtavarre 18 Jul 1899 Haglund & KaII- strom (UPS!). - Sor-Trondelag, Oppdal, Kongsvoll 11 Aug 1884 Bryhn (UPS!); Mt. Knutsho 7 Aug 1863 Fries (UPS!), 19 Jul 1870 Wickbom (UPS!), 10 Aug 1888 Blytt (Blytt & Rostrup 1891 as Trochila juncicola, O!), 20 Jul 1894 Dyring (UPS!), July 1898 Carling (UPS!). - Sweden: Torne Lappmark, Kiruna (formerly Karesuando) Mt. Paltsa reg. alp. 27 Jul 1919 Alm & Tengwall (UPS!), E slope of S summit 1100 m s.m. 24 Jul 1933 Smith (UPS!), N slope of N part 1100 m s.m. 2 Aug 1939 Smith (type, UPS!); (formerly Jukkasjarvi) Mt. Kartetjikka 1400 and 1300 m s.m. 15 Aug 1925 and 9 Aug 1927 Smith (UPS!). - Lule Lappmark, Jokkmokk Mt. Jille 17 Aug 1915 Tengwall (UPS!). - Canada: Distr. of Franklin, Ellif Ringnes I s l . 3 finds (Groves & Elliot 1971 as H . pusilla, DAOM). Acknowledgements - During the four decennia that I have been interested in the small emergent Discomycetes on Junca- ceae my studies of them have been promoted by too many per- sons to be enumerated here, but I am most grateful to them all. A few must however be specially mentioned. Professor R. San- tesson scrutinized much of the material of Juncus in UPS in search for these fungi. Mrs Kerstin and Dr L. Holm brought much important material, and Dr Holm checked the Latin di- agnoses and critically read the manuscript. With Dr B. Hein, working upon a thorough revision of the whole genus, I have had a most instructive and stimulating correspondence, and when the present paper was approaching its definite shape, he most generously sent me a preliminary draft of his manuscript. Mrs Ulla-Britt Sahlstrom performed skilful photographic work, and Mrs Ulla Hedenquist patiently and carefully typed the manuscript. My thanks are also due to the directors and staffs of the herbaria which have been consulted: C, DAOM, GZU, H, K, 0, PAD, S , UME, UPS, ZT. References Ainsworth, C.G. 1971. Ainsworth & Bisby’s Dictionary of the Fungi (ed. 6). - Commonwealth Mycological Inst., Kew. Blytt, A., & Rostrup, E. 1891. Bidrag ti1 Kundskaben om Nor- ges Soparter 2. Ascomyceter fra Dovre. - Kristiania Vid.- Selsk. Forh. 1891: 9. Clements, F. E. 1909. The genera of Fungi. - Minneapolis. - & Shear, C. L. 1931. The genera of Fungi - New York. Corda, A. C. I . 1838. Icones fungorum 2. - Calve, Prague. Dahlgren, R. M. T. & Clifford, H. 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