Stray Studies in the Coronophorales (Pyrenomycetes) 4-8 - ASCOfrance

Studies in the Coronophorales 4-8 291Through the courtesy of Dr. Sivanesan I have had access to a fragment of IMI 46511 (a)(unfortunately devoid of asci) and slides of IMI 165 094. The ascocarps resemble in shape thoseof Thaxteria and lack certainly a functioning ostiolum, although the apical part of theascocarpic locule possesses thick conspicuous periphyses. The rest of the locule is lined by acontinuous palisade of asci at very different stages. No apical apparatus has so far beenobserved, but richer material is needed. The peridium shows tendencies to being slightlyareolate in the same manner as in sundry Lasiosphaeriae and Cercophorae. The cell wallsshow a kind of "Munk pores", but these are smaller than usual and the surrounding thickenedring less prominent.It should also be observed that the inclusion amongst the Coronophoralesspecies of Bertia (B. moriformis)of the typedoes not mean that other species placed in thatgenus necessarily are of the same affinity. Only three have been examined, viz. B.puttemansiiP. Henn. (=Thaxteria archeri, see above), B. italica Sacc. & Speg. andB. (Bertiella) macrospora Sacc. and they were all found to be alien elements, thethird even bitunicate. The second was synonymized by Mtiller & Arx (1962: 818)with Calyculosphaeria macrospora Fitzp., but this is a true Nitschkia (N.gigantospora).We shall now compare various significant features in the species studied by me. Itwill then gradually emerge that Fitzpatrick's Nitschkieaeafter some remodellingstands out as a group of inter se closely related species, whose "pattern of variation"suggests a multidimensional letticework of crossing lines (comp. Munk 1962),whereas other Coronophoralestake more isolated positions.Mode of growth and ecology. All Coronophoralesgrow on bark and/or wood, butthe principal habitat of one (N. parasitons) is stromata of Nectria cinnabarina and/orits conidial state Tubercularia vulgaris (see e.g. Schweinitz 1832; Tulasne1865=1931; Petrak 1927:362; Chesters 1939; Mason 1940; Muller & Arx 1955:366;Môller 1958). Stromata may at the same time bear well-grown ascocarps of both theNectria and the Nitschkia. Sometimes, when the Nitschkia is unusually vigourous, itmay also form patches of closely crowded ascocarps—like those of the otherNitschkiae—onthe surrounding bark. This species seems thus to be ± parasitic.Desmazières' statement (1860 n. 780) that his specimens (on Robinia pseudacacia) "sontquelquefois mélés avec le Nectria coccinea, et même, mais plus rarement avec le N. Cinnabarina"rests on a clear misunderstanding of the former species, for he continues "cesespèces reposent sur le Tubercularia confluens, Pers. var. Acaciae, Fr. qui leur sert destroma". His "N. coccinea" must thus have been N. cinnabarina too.The remaining species seem to be saprophytic, but it may be doubted if any is a"saprophyte of the first incidence" (sensu Munk), and indications of hypersaprophytismare numerous. Such are e.g. the frequent association of N. confertulaold stromata of Hypoxylon rubiginosum observed in Britain (Chesters I.e.; MasonI.e.), U.S.A. and South Africa (Miller 1942:259) as well as in Ghana (IMI).with20-753873 Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 29320 /am in diam.) (Fig. 2 j). In Ac. tristis they are especially numerous round the basesof the ascocarps and form there distinct "crowns". In the other species they arepresent also on the very ascocarps (see below).Ascocarps. Unfortunately, very little is known about the ontogeny of the Coronophoraleanascocarps, and practically nothing about those of the Nitschkieae. Bertiamoriformis has been studied in some detail by Luc (1952) and Parguey-Leduc(1966: 19-26), and Coronophora gregaria (Lib.) Fuck, by the latter authoress (I.e.pp. 26-30). Scattered notes are contributed by e.g. Petrak (1921: 182-184, Coronophora)and Münk (1953:34-35, 96-100; 1957:288-295).It is evident that the development does not follow the "classical" ascohymenialscheme (i.e. the "Xylaria Type" of Luttrell 1951), the interior of primordia andyoung ascocarps being filled by a hyaline thin-walled tissue (thus corresponding withLuttrell's "Diaporthe Type"). The asci are formed within this tissue, which changesin various ways and successively dissolves. A central "locule" appears, containingthe asci and lined by hyaline compressed cells. The asci are often seen to be attachedto a basal large-celled placenta-like cushion or to some few smaller lateral ones.Such a basal "coussinet ascogene" was described and depicted in Bertia by Luc(I.e.) and Müller & Arx (1962:817). It has also been observed by me, but Parguey-Leduc (I.e.) could not find it. I have seen similar structures in several other species,and Arx & Müller (1954) show them in N. acanthostroma (fig. 116) and N. similis(fig. 117), Müller & Arx (1955) in N. confertula, Sivanesan (1974:38, fig. 2a) in N.leonensis.The basic shape of the mature ascocarp in Fitzpatrick's Nitschkieae (well shownin his excellent photographs) is turbinate or kettledrum-like and (especially when dry)collapsed into cup-shape. There is no true (perforated) ostiolum but mostly a distinctapical papilla. The surface is finely to coarsely tuberculate. Enclosed in the upperpart is the ± flattened locule. N. broomeiana (=Fracchiaea heterogenea) is extremeby its unusually robust and firm ascocarps, which only rarely collapse.In some species the ascocarps have a voluminous base, so high that they lookstipitate and their heights exceed their widths, but in others it is almost absent, thedry ascocarps looking as sessile apothecia. The first case may be exemplified by N.calyculus and N. parasitans, the second by N. collapsa, N. pezizoidea and N.tetraspora.The size of the adult ascocarps varies within rather wide limits, but is ratherconstant in the individual species. The smallest ascocarps (ca. 0.2-0.25 mm across)are those of N. parasitans, the usual size is 0.3-0.5 mm, and in some species (e.g. N.collapsa and N. broomeiana) they may reach 0.8 mm.The peridium is a coriaceous (not carbonized) pseudoparenchyma of in the outerparts brown-walled, rather large cells. When there is a stipe-like base, its internalcells are prolonged vertically and arranged in slightly divergent rows. The external20-753873 Svensk Bot. Tidskr. 69 (1975)

294 J. A. Nannfeldttuberculations are formed by aggregations of ± globose cells with walls thicker anddarker, especially peripherally. In a few species some peripheral cells run out intopointed, spine-like processes, which in N. brevispina have an average length of ca.10 ¿im (Munk 1957:185, fig. 67) and in N. broomeiana, where they often are forkedand/or provided with secondary "barbs", may reach a length of 25 /urn or more. InAc. argentinensis, Ac. foveolata and Ac. pulchella the ascocarps are armed with arestricted number of easily broken-off bristles agreeing with those of the subiculum(see above).The basic shape of the ascocarps in HöhnePs Coronophoreen (excl.Coronophorella)is subglobose or broadly ellipsoid, without ostiolum and apical papilla, shrivellingirregularly on drying and only rarely distinctly tuberculate. The ascocarps arerelatively large, in some species (e.g. Coronophora gregaria) even very large,reaching almost 2 mm in diameter. The structure of the peridium agrees with that inthe Nitschkieae.The ascocarpic wall is only slightly thicker basally.In the two genera later added to the Coronophorales, Bertia (moriformis) andGaillardiella (pezizoides)the ascocarps possess well developed basal parts. In theformer the ascocarps are subcylindrical, large and high (up to 0.7x1 mm) and verycoarsely tuberculate (whence the specific epithet). They have no apical papilla anddo not collapse on drying. The apex is unrecognizable externally because of thetuberculations, but thin, exactly median sections show a small "plug" with specialanatomy simulating an ostiolum (Figs, la, b). Due to its minuteness the "plug" wasmissed by Luc, Munk, and Parguey-Leduc, but it was illustrated and described byMüller & Arx (1962:816-818).The ascocarps of G. pezizoides are also rather large (up to l.lxl.l mm) andsubcylindrical but collapse cupulately on drying. The extant material (in FH) is nowso poor that we have in the main to rely upon the detailed studies by Petrak (1953a)and Müller & Arx (1952:818-820). The latter authors, who also draw a section,recognized it as Coronophoralean. It approaches Nitschkia by the collapsingascocarps, but deviates markedly by their size and, especially, by the strong circularthickening of the peridium round the "cup"."Munk pores" (Nannfeldt 1975:51) have been observed in all members studied byme, thus also in the large-fruited (true) Coronophora gregaria, where Munk himselffailed to see them, but their obviousness is very variable, depending both upon thespecies and the stage.The pores are ca. 1 p.m in diam. and surrounded by a ring-shaped thickening of thewall, which makes them look like miniatures of the ring pores of the conifers (Fig.2e). Their number varies with the species. As a rule the common wall between twocells shows only one pore, but on some occasions a higher number seems out ofdoubt.By the way it may be mentioned that Patouillard seems to have been the first to observe thesepores. He drew them carefully in a pencil sketch accompanying the type specimen of hisSvensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 295Gaillardiella pezizoides,did not appear in print.where they are both numerous and conspicuous, but his observationTowards the locule the peridial cells become flattened and their walls thinner andpaler. The cells lining the locule are hyaline-walled and almost compressed. Apartfrom the ± strongly developed base the peridium is of almost uniform thickness,except that, in some species without a "Quellkôrper' ' (see below), the brown layer isdistinctly excavated apically, the hyaline tissue intruding into the papilla as a domeor flat cone, e.g. N. broomeiana (Figs. 2a, b), N. collapsa (Fig. 2h), N. grevillii(Figs. 2c, d; further ill. Muller & Arx 1962:815 = 1973:92), and N. tetraspora. IfPetrak's description and drawing (1952d) ofN. velutina are correct, the brown layerhas (almost) disappeared apically and no papilla is visible in it. The hyaline tissueshows always special structures below the papilla (see below), and due to theirfirmness the papilla persists or becomes even more prominent after drying.Much has been written about the mode of spore discharge in the Coronophorales,but little is in fact known about what happens in nature. Fuckel (1871:324) describeshis Coronophora macrosperma (a true Coronophora fide Hôhnel 1918: 137) with"ascis ut massa hyalina expulsis", and Patouillard & Gaillard (1889:107) remarkabout their N. multiasca:—"Le contenu des périthèces s'échappe sous forme d'unemasse blanche". Boudier (1910a: pl. 574) depicts his Trichosphaeria vagans (=N.collapsa) with slime drops crowning some ascocarps and comments (19106/ 340):"les uns émittent des globules de spores et d'autres sont dépressés circulairementprès de l'ostiole après leur sortie". My sole personal observation is that, in a freshsample of N. parasitans (Holm 175a), a few minutes after thorough wetting somemature ascocarps ruptured on one side by an irregular subequatorial slit and openedas by a hinge, thus exposing the slimy spore mass. In the same way behaved also anold, very well developed herbarium sample (Romell 15834, S).The textbook descriptions are taken from Hôhnel's detailed accounts, but theseare based on his experiments with herbarium material, placing ascocarps in waterand studying them in the microscope. They must thus be taken cum grano salis. Hefound that in Coronophora annexa (Nke) Fuck. (1906:662-663) and N.chaetomioides (1910:1505-1507) mature ascocarps swelled, their peridium burstapically, a hyaline sack protruded and emptied its contents of spores embedded inslime through an apical, gelatinized pore. Afterwards a compact subcylindrical bodywith transversely elongated cells with strongly gelatinized walls became visible. Heinterpreted this body as a rest of the contracted sack. He studied later(1918:136-137) richer material of the former species and found his earlier interpretationto have been erroneous, for the gelatinized body was present also in unopenedascocarps, hanging down from the apex of the locule. These bodies were nowconsidered as "Quellkôrper, die im Augenblicke des Aufreissens der Perithezien sichplôtzlich stark vergrôssern, das Perithezium zum grossen Teil ausfiillen und20-753873 Svensk Bot. Tidskr. 69 (19

296 J. A. NannfeldtFig. I. (a. b) Bertia moriformis 3.x. 1948 Lundell UPS) median sections of ascocarps showing the ostiolumlikeapical structure of the pendium (ca. 200x, ca. 450x). — (c) Nitschkia chaetomioides (IMI 40291 r)median section with "Quellkorper" (ca. 160x). _ (d) N. calUsta (Sturgis UPS) top of "QueUkorper" withperiphysis-like hyphae (ca. 400x). _ (e,f)N. confertula (Curtis UPS) eccentric sections, e showing theenlarged base of a "Quellkorper",/showing a curved "Quellkorper" in cross-section (ca. 160x). — (g) N.acanthostroma (IMI 53244) median section (ca. 160x). — (/,) N. acanthostroma (Moller COI) subicularhyphae (ca. 250 x).Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 297Fig. 2. (a, b) Nitschkia broomeiana (1888 Bresadola UPS and IMI 139 074) median sections of ascocarps, anolder stage in b (ca. 150x). — (c, d) N. grevillii (IMI 68040) median sections (ca. 40x, ca. 150x). — (e)Acanthonitschkea foveolata (Malme S) "Munk pores" (ca. 400x). — (f, g) N. parasitans (Munk UPS andIMI 9901) median sections, an older stage in g (ca. 250x, ca. 150x). — (h) N. collapsa (Romell 15823 S) and(/) N. florida (Hb. Fitzp. 1896 FH) median sections (ca. 150x). — (/) Ac. foveolata (Wright LPS) subicularbristle (ca. 200x).hierdurch den Nukleus aussleudern". The same structure was independently describedand depicted by Werdermann (1923) in N. acanthostroma.When revising Fracchiaea Fitzpatrick (1924) took for granted that "Quellkörper"were structures unique (but not obligatory) to Höhnel's "Coronophoreen" and alien20-753873 Svensk Bot. Tidskr. 69 (1975)

298 J. A. Nannfeldtto his own Nitschkieae, and so he transferred three alleged Fracchiaeae with"Quellkörper" and cupulate ascocarps to Höhnel's group. In the present paperthese species are put back into Fitzpatrick's group and included in Nitschkia.year before, when establishing his Nitschkieae,TheFitzpatrick (1923) had evidently notbecome observant of this structure, for in one of his own still preserved slides (FH)of Ac. macrobarbata (i.e. Ac. foveolata) a large "Quellkörper" is readily seen.It must be presumed that the "Quellkörper" play an important rôle in the sporedischarge or at least have played such a rôle during the evolution of the Coronophorales,but how they function is not known.Such ± prolonged subcylindrical to inverted-conical "Quellkörper", which mayeven reach almost down to the bottom of the locule, occur in the followingNitschkieae:Ac. foveolata (ca. 300 fim long)Ac. pulchella (ca. 200 fini)N. acanthostroma (ca. 250 /xm; Fig. \g, further ill. Werdermann 1923, Arx & Müller1954:379 (inaccurate)N. affinis (rather short?)"Fracchiaea australis" (ca. 200 /im; ill. Fitzpatrick 1924)N. callista (rather short?; Fig. Id)N. chaetomioides (ca. 300 /xm; Fig. lc)N. confertula (rather short; Figs, le,/, further ill. Müller & Arx 1955 (inaccurate)N. leonensis (ca. 250 ju.ni; ill. Sivanesan 1974)N. similis (ca. 200 /im; ill. Arx & Müller 1954)N. uniseriata (ca. 120 /im).The "Quellkörper" consist of concentrically and transversely orientated, firmlyconglutinated cells with very thick, hyaline, strongly refractive walls and narrowlumina (Figs, lc, d,f). The parabolically roùnded apex shows a fringe of longitudinalhyphae. In N. callista these hyphae have thick walls and narrow lumina and curveparabolically inwards. In N. leonensis they are thin-walled with larger lumina (thusresembling normal periphyses), rather straight and convergent as to circumscribe acone. N. chaetomioidesis ± intermediate with curved and rather thick-walled hyphaebut still ample lumina.In other Nitschkieaethe "Quellkörper" is replaced by a low perforated "cushion".Höhnel (1918:138) found that in Fracchiaea heterogenea (=N. broomeiana)the hyaline layer of the peridium thickens apically (from ca. 20 /xm to ca. 70 /xm),"ist hier rundlich durchbrochen und löst sich hier in einen Kranz von Periphysenauf. Diese innere hyaline Schicht hat daher ein typisches Ostiolum". Such "cushions",which may be interpreted as either incipient or reduced "Quellkörper", haveso far been observed in:Gaillardiella pezizoides (ill. Müller & Arx 1962)N. brevispinaN. broomeiana (Figs. 2a, b)Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 299N. collapsa (Fig. 2 h)N. cupularis s. orig.N. floridana (Fig. 2 i)N. grevillii (Figs. 2c, d, further ill. Müller & Arx 1962, 1973)N. parasitans (Figs. 2f, g)N. velutina (ill. Petrak 1952d).The "cushions" of N. parasitans(Figs. 2f, g) with its peerlessly small ascocarpsare of special interest, for their hyphae are unusually periphysis-like and leave arelatively large, upwards tapering, empty canal reaching the brown, outer peridiallayer, which shows no sign of an opening, not even of an excavation.With regard to Höhnel's "Coronophoreen" it may suffice here to note thatCryptosphaerella is marked by the presence of "Quellkörper" and Coronophoras.str. by their absence.When an ascocarp of the Coronophoralesis crushed under the cover-slip thelocule surrounded by a hyaline membrane may sometimes protrude as a closed sack.I have not been able to establish, whether this behavior is peculiar to certain speciesonly and/or to certain stages. Nor do we know, whether this behavior has anycounterpart in nature or is a mere artifact (comp. Höhnel's observations citedabove). Cells and cell-rows of the hyaline lining layer loosen often in squash-mountsand represent certainly the paraphyses and pseudoparaphyses of various previousdescriptions. The readiness with which the lining layer loosens from the peridiumstrongly favours a supposition that, ontogenetically, this layer is part of the locule,i.e. of the (perithecial) centrum sensu Wehmeyer (1926) et Luttreil (1951), not of theperidium.In my opinion, the above comparative studies of the ascocarps show convincinglythat the Coronophorales (after the removal of Thaxteria, Rostrocoronophora,Spinulosphaeriaand some odd species) form a most natural unit. The close agreementin structure and texture and such striking features as "Munk pores" and"Quellkörper" make a double origin such as suggested by Parguey-Leduc (1966)most improbable, to say the least. She may have been influenced in part byChadefaud's (1960:588-589) confused conception of the Coronophoralesas embracingalso such fungi as Calosphaeria princeps Tul. and Enchnoa injemails (Kze exFr.) Sacc.With regard to the shape and structure of the ascocarps Fitzpatrick'sform a very homogeneous group, to which also FracchiaeaNitschkieaebelongs and Gaillardiellashows close affinity. Höhnel's Coronophoreen (i.e. Coronophora s.lat.) form asecond very homogeneous, smaller group, whereas Bertia alone stands out as a thirdgroup, but the differences are not profound enough to motivate different families.Asci. As already mentioned, the ontogeny of the asci is not sufficiently known. Mostasci of a locule are in about the same stage, and young asci are relatively rarely seen,suggesting that the early stages pass rather rapidly.20-753873 Svensk Bot. Tidskr. 69 (19

300 J. A. Nannfeldt4\*n»e1 ,iNlgM^ ?w *4 * W J * '?E H F vFig. 3. (a, b) Coronophora annexa (Krieg., F. sax. 968) asci (ca. 725x,ca. 900x). —(c-/)Nitschkia collapsa(c, d: 28.iii.1908 Lind C; c in Lactic Blue, d in glycerine with Congo Red) asci, the apical cylinder clearlyseen in d (ca. 1600x); (e, f: Romell 15823 S) spores (ca. 900x). — (g) C. gregaria (Petr.. Myc. gen. 1567)ascus (ca. 725x). — (h)N. callista (Rel. Farl. 706), (i)N. uniseriata (typus), and (j)N. affinis (typus)asci (ca.900x). — (k) N. broomeiana (Rav., F. carol. 4: 57) asci (ca. 725x). — (I, m) N. confertula (Curtis UPS andIMI 144 570) asci (ca. 725 x). — («) N. gigantospora (typus), (o) N. calyculus (typus), (p) Acanthonitschkeafoveolata (Malme S), and (q)Ac. argentinensis (IMI 38113a) spores (ca. 900x).Svensk Bot. Tidskr. 69

Studies in the Coronophorales 4-8 301Generally speaking, the asci are stipitate, rather thin-walled and obviously unitunicate.As a rule the wall deliquesces rather early. The apical apparatus has lostits gun-function and is abortive.In Coronophora the shape of the ascus (Figs. 3 a, b, g) is most characteristic(comp. e.g. Parguey-Leduc 1966: 26-30) and repeatedly illustrated. The wall of themature ascus is relatively firm and slightly thickened in the apical part, which iscylindrical to conical and obtusely rounded. Downwards, often delimited by a slightconstriction of the lumen (a "bourrelet sous-apical"), follows the main body of theascus as a bulging sack, downwards ± abruptly constricted into a very long, almostfiliform stipe. In a subadult stage the "bourrelet" is much stronger, and so thesubapical chamber of the spore-stuffed lumen looks like a stalked head "crowning"the body of the ascus as depicted by the author of the genus (Fuckel 1870 tab. vi, fig.16) and commemorated in the generic name.In the Nitschkieae the asci are more varied. Those of N. collapsa are picturedwith an apical wall-thickening and a cylindrical apical apparatus by Boudier (1910api. 574) and Lind (1913 tab. iii, fig. 31). Chenantais (1918:71-72, fig. 5b) depicts twoyoung asci and describes them as follows: "A l'état jeune, ils sont surmontés d'uncylindre hyalin pourvu au centre d'une dépression en entonnoir fermée par unanneau réfringent qui se présente en coupe optique sous l'apparence de deuxguttules, analogues à celles des Diaporthe, Melanconis, Laestadia etc.". Fitzpatrick(1923) notes an apical wall-thickening in the same species as well as in TV. cupularisand N. grevillii. Arx & Millier (1964:815=1973:92) describe and depict it in thelast-mentioned species.These structures are only faintly seen in lactophenol (and in Lactic Blue) but wellshown in water, which certainly was the medium used by the early authors.Although the structures themselves do not take Congo Red, this stain makes themstand out more clearly. Janus Green as a stain has also sometimes proved to beuseful as well as glycerine as a medium. In N. collapsa the apical thickening is justdiscernible in Lactic Blue (Fig. 3 c), but an ascus in the right stage shows in glycerineand Congo Red an apical perforated cylinder, up to 2 pm high and about as broad(Fig. 3d). In N. cupularis a distinct wall-thickening is seen with Congo Red. Theapex is often truncate and may even show a deep hemispherical invagination.It seems likely that, at least during some (short) stage, the ascus wall of most (all?)species is thickened apically and has a "bourrelet sous-apical". Thus e.g., in N.floridana the tip may show a slight thickening and be truncate and even shallowlyconcave. In N. brevispina, N. tetraspora, Ac. foveolata, and Ac. tristis asci havebeen seen with the tips constricted into short, truncate cylinders with slightlythickened wall, and in the first also an apical apparatus in the shape of a low,perforated cylinder taking Janus Green. In N. parasitans a distinct wall-thickeninghas been observed and, occasionally, also an abortive ring has glimpsed as twoindistinct points (comp. Chadefaud 1960: 587 fig. 444: 2 as N. cupularis).20-753873 Svensk Bot. Tidskr. 69 (19

302 J. A. NannfeldtThe shape of the (mature) ascus is as a rule ± clavate to fusiform with the sporesdisorderly in several rows. In a few species they are ± cylindrical, thus in N. affinis(Fig. 3j), where they are rather truncate, and in N. uniseriata (Fig. 3i), where theeight spores are uniseriate and often transversely orientated. Tendencies to transversespore orientation are pronounced in both Coronophora and the Nitschkieae,thus in Coronophora annexa (Figs. 3a, b) with its polysporous asci and allantoidspores, and in an early stage in Ac. foveolata, when the eight spores are alsouniseriate.Number of spores. True polyspory (sensu Martens 1937) is common within theCoronophorales, and the number of spores per ascus is then often about 32.All species of Coronophora s.lat. are polysporous. To judge from the literature(e.g. Hohnel 1906, 1907; Munk 1957) the number of spores should normally be about32, but one (anonymous) species illustrated by Munk (I.e. fig. 111c) seems topossess only about 16 spores, and C. annexa as illustrated by Miller (1949:100 fig.31) shows much more than 32. My own observations on the latter species (Krieger,F. sax. 968) indicate that the number often approaches 64, but that some asci haveonly about 32. About 64 seems to be the normal number in C. gregaria (severalcollections).In Bertia moriformis the asci are 8-sporous, and so they are in the majority of theNitschkieae, but six species have about 32 spores per ascus, viz. Ac. pulchella, N.affinis, N. callista, N. leonensis, N. multiasca, and N. similis. Deviations from evenmultiples are not rare and in some species rather the rule. Thus in N. affinis and N.leonensis numbers about 25 seem to be the most common, but in the latter speciesI have observed numbers up to about 32 and down to about 16.N. broomeiana offers an extreme case not only by the high number of spores(>200) per ascus but also by their extraordinary and most characteristic arrangement,excellently described by Fitzpatrick (1924:108): "in several longitudinal rows,each spore lying obliquely with its lower end towards the ascus wall and its upperend towards the interior". I know of only one similar arrangement, and that waskindly pointed out to me by Dr. N. Lundqvist, viz. the oblique orientation of thespores in the young ascus of Podospora setosa (Wint.) Niessl (comp. Lundqvist 1972pi. 28 fig. d).My new African Nitschkia deviates in the opposite direction. Though 8 spores areformed in the ascus, only four of them reach maturity (hence the epithet tetraspora),become fusiform and get a coloured wall. The remaining four (as a rule the basalones) remain as acicular to narrowly fusiform, very thin-walled bodies withhomogeneous plasma and sometimes a spurious septum. They are best seen just afterthe deliquescence of the ascus wall, as they deliquesce later soon themselves.Spore size, shape, septation and wall. The spores are mostly relatively small, suballantoidto boomerang-shaped, until late or permanently continuous with thin,Svensk Bot. Tidskr. 69

Studies in the Coronophorales 4-8 303smooth, long or permanently hyaline walls, without germ pores or germ slits,without gelatinous sheaths, and at least in some stages, with distinct drops ordroplets, ± constant in number.Strikingly divergent are the spores of Bertia moriformis. They have the usualappearance but are of another magnitude (30-50x5-6 /xm), become normally 1-3-septate but may occasionally get up to 7 septa (Hawley 1923:223). About as large arethe spores of N. gigantospora (Fig. 3 n) and N. macrospora.Suballantoid, permanently hyaline and sometimes eventually 1-septate spores (as arule 8-12x1-2 pm) characterize almost all species with polysporous asci (both inCoronophora and in the Nitschkieae) but also some with 8-sporous, e.g. N. parasitans.The allantoid spores of Ac. argentinensis (Fig. 3 q) are extreme both in theirminuteness (ca. 5-8x2 pm) and in their curvature (often semicircular or horseshoelike).In some species the subcylindrical spores are only slightly curved to almoststraight, e.g. in N. calyculus (Fig. 3 o), N. grevillii and N. cupularis.A peculiarity, on which two mono typical genera have been based, is the apicalappendiculation of the spores in N. velutina {Biciliospora; not seen by me) and N.similis (Scortechiniella). The spores of the latter show also another, hitherto unnoticedpeculiarity, unique within the Coronophorales, viz. a spiral twisting (up to90°).In N. confertula (Figs. 3 /, m) the spores are ± reniform, inequilaterally ellipsoidor ovoid, get early coloured (smoky-grey) walls but remain unseptated. In N. collapsa(Figs. 3 e, f) and N. tetraspora (Figs. 4 b, c) the spores are straight, becomeearly fusiform and distinctly 1-septated, and get early coloured walls.Only one species (N. uniseriata) has an ornamented wall and is so an analogon toLasiosphaeria punctata Munk (1957:114-115) in that genus. At a certain stage, whenstill within the ascus, the spores of N. uniseriata possess distinct cyanophilous wartson the hyaline wall (Fig. 3/), but in fully mature spores with coloured wall the wartshave been reduced to a barely visible roughness.Taxonomic conclusions. The above comparative studies on asci and spores reveal asurprisingly wide variation, but this does in no way weaken the conclusion of thehomogeneity of the Coronophorales, won by the morphological studies on theascocarps.The peculiar ascus shape, the polyspory, and the spore shape can be added to thecharacteristics of Coronophora. Bertia keeps its isolated position. But the variablefeatures within the Nitschkieae do not show any correlation inter se nor withfeatures in the ascocarp morphology and disclose no evolutionary trends. On theother hand, they demonstrate convincingly the futility of genera in this group, basedexclusively on such characters, and this invites to a complete merging of all theNitschkieae into one genus. However, four species stand out by the addition of apeculiar morphological structure, viz. the long, stiff, dark bristles on the subiculumSvenskBot. Tutskr. 69 (1975)

304 J. A. Nannfeldtand, in three of them, also on the ascocarps. These bristles are exactly alike in allspecies, and their monophyletical origin can hardly be questioned. Even if thespecies are not homogeneous as to "Quellkorper" and spore number I find itmotivated and preferable to treat them as a separate genusdistinct from the bristle-lessNitschkia.(Acanthonitschkea)My conclusion is thus that all Coronophoralean fungi so far known should betreated as one family (Nitschkiaceae) with Coronophora, Bertia, Gaillardiella,Nitschkia and Acanthonitschkea as the genera.The establishing of the correct name for the combined family offers its problem. To thosewho accept "Familie der Coronophoreen" (Hohnel 1907) as a valid name, Coronophoraceaeby priority becomes the correct name. But to those who—like myself—consider the Latinending as a sine qua non, Coronophoraceae and Nitschkiaceae are of the same age (Nannfeldt1932), and so we have to follow Miller (1949:121) in using Nitschkiaceae, for as far as I amaware he was the first to combine the two families.In this family, and especially in the "Nitschkieae",evolution and speciation haveworked with "Quellkorper", ascus structures and spore shape, whereas ascocarpshape and peridium structure have remained fairly constant. The family affords thusa striking contrast to another aberrant, most characteristic and homogeneous groupof true Pyrenomycetes recently studied by me, viz. Boliniaceae with Camarops asthe sole genus (Nannfeldt 1972). In that family asci and spores are practicallyidentical in all species, whereas perithecial shape and stromatal size and shape showenormous differences, the stromatal shape varying from almost valsoid to xylarioid,but the variation in stromatal structure is still kept within narrow limits.It remains to discuss the taxonomic position of the Coronophorales(=Nitschkiaceae). To-day considerably more is known about the evolutionarytrends within the Ascomycetes than it was well 40 years ago. One of the salientfeatures of the Coronophorales, the cleistocarpy, is now known to have evolvedrepeatedly and is not eo ipso a sign of high taxonomic rank (comp. e.g. Cain1972:3-5; Miiller & Arx 1973:96). The abortive apical ascus apparatus is also anobviously polyphyletic phenomenon and in our case a natural consequence of thecleistocarpy. The recognition of the "Diaporthe Type Centrum" (Luttrell) makes itpossible to connect the Coronophoralean locule with more "normal" ascohymenialtypes.The features of our group are thus no longer so strange as originally thought, andits place among the true (ascohymenial and unitunicate) Pyrenomycetes seemsfirmly established. Here it should be understood that, in my opinion, there are noconvincing facts in favour of the hypothesis by Chadefaud and his school thatunitunicate asci have evolved polyphyletically from bitunicate. Recent investigationson the fine structure of the ascus walls make such repeated transitions evenmore unlikely (comp. e.g. Griffith 1973).Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 305Next question is then, in what direction the affinities of our group might be sought.Carroll & Munk (1964: 96-97) suggested "that the natural affinities of the Coronophoralesare very close to the genus Lasiosphaeria", the crucial species being theirnew L. noonae-daniae. Its ascocarps are ostiolate, and its spores germinate byphialides, but otherwise it exhibits a number of Coronophoralean features: tuberculateperidium, radiating vertical cell-rows in the bases of the ascocarps, "Munkpores" (also in L. tuberculosa Carroll & Munk), stipitate asci without any apicalapparatus. An additional indicium is the cyanophilous network of longitudinal,anastomosing cytoplasmic ribs on the inside of the wall in emptied asci. A possibleconnection in about the same direction (with Bombardiella and Bombardia) wassurmised already by Theissen (1917).With Carroll & Munk's suggestion as the working hypothesis we shall nowcontinue the comparison between the two groups. As the Coronophoralean asci as arule deliquesce early, cyanophilous ribs have not been sought for. "Munk pores"have been found in all Coronophorales and also in Thaxteria and Spinulosphaeria(? comp. above), two genera above transferred to Lasiosphaeriaceae. Divergingvertical cell rows in the basal peridium occur in all Coronophorales with "stipitate"ascocarps, and the same trend has been found by Parguey-Leduc (1967:373) inanother ally ("L. ovina" =Bombardia (?) sp. fide Lundqvist 1972:77). It occurs alsoin Thaxteria and Spinulosphaeria. The peridium structure is very uniform in ourgroup, highly diverse in Lasiosphaeria, including such of Coronophoralean type.The "Quellkórper" are unique to our group but the "apical cushions" of the loculesof some species resemble closely the periphysate "inner" ostiola in Lasiosphaeria(comp. Parguey-Leduc 1973:75 fig. 4D), and so do the fringed tips of the"Quellkórper". The simplification of the ascus structures in our group is well on theline with similar trends within Lasiosphaeria and allies. The spore germination byphialides is a peculiarity within the Lasiosphaeriaceae, where the spores as a ruleare large and 8 per ascus but extremely varied as to shape, septation and colour. Afew Coronophorales have large spores, but the majority medium-sized to small. Truepolyspory (combined with allantoid spores) is widespread in our group.Hypersaprophytism is common in both groups.My final conclusion is thus that there are no longer sufficient reasons for keepingour group as an order of its own but that it still deserves the rank of an independentfamily (Nitschkiaceae), which should be regarded as a highly specialized "satellite"group derived from Lasiosphaeriaceae sensu Lundqv. It will presumably—like thatfamily—be found to have its place in a remodelled order Sordariales (comp. Lundqvist1972:55-56).5. The Species of NitschkiaThe considerations in the preceding Chapter result in a wholesale reduction of thenumber of accepted genera and in the following long nomenclátor for Nitschkia,SvenskBot. Tidskr. 69 (1975)

306 J. A. Nannfeldtembracing no less than 16 generic names based on its well 20 species. (In thenomenclators for the individual species appear a total of 20 additional generic namesthat Nitschkiae have been described under or combined with and as synonyms 20specific and infraspecific epithets!)Somebody may consider it a mere waste of paper to publish these nomenclators insuch detail, but the bitter words on the polypores by my late friend M. A. Donk areof general applicability: "During these activities I encountered a complaint datingfrom as long as 150 years ago that revealed that a failing common to many a modernauthor is of long standing: 'The practice of accurately examining the synonyms,instead of copying them from other authors, has been too long neglected' (Purton,App. Midi. Fl. 335. 1821). The failing has culminated in a number of errors andpuzzles in references in some of the much used European publications that is reallystupendous" (Donk 1974: 5-6).Nitschkia Otth in Fuck., Symb. mycol. p. 165 (1870; nom. nud.); ex Karst., Mycol.fenn. 2: 13 (1873). — Lectotypus: N.fuckelii Nke=A f . cupularis sensu Karst., i.e. N.parasitans.[Cyathisphaera Dum., Comment, bot. p. 87 (1822) p.p. (vide Nannfeldt 1975:60-61).]Coelosphaeria Sacc., Atti Soc. Ven.-Trent. Sei. Nat. 2(2): 163 (xii.1873) — Nom. nov. proNitschkia, diatomearum generis Nitzschiae Hassall causa; nom. nud. et superfl. (cp. NannfeldtI.e.).Fracchiaea Sacc., ibid, (xii.1873). — Typus: Fr. heterogenea Sacc., i.e. N. broomeiana.Echusias Hazsl., Verh. zool.-bot. Ges. Wien 23:367 (xii.1873). — Typus: E. Vitis "(Schulzer)"Hazsl. [non Gibbera Vitis Schulzer], i.e. N. broomeiana. — Syn. fide Höhnel 1920:130-131.Scortechinia Sacc. in Sacc. & Berl., Atti R. Ist. Venet. Sei. Lett. Art. 6(3:4): 713 (1885). —Typus: Sphaeria acanthostroma Mont.Tympanopsis Starb., Bih. K. Sv. Vet.-Akad. Handl. 19(3:2):24(1894). — Typus: Sphaeriaeuomphala Berk. & Curt., i.e. N. confertula.Winterella Berl., Icon, fung. 1(3):94 (1892) [non Winterella O. Kze (1891) nec WinterellaSacc. (1899)]. — Typus: Winteria tuberculífera ("tuberculigera") Ell. & Ev., i.e. N. grevillii.Winterina Sacc., emend. Sacc. & Syd. in Sacc., Syll. fung. 14:589 (1899) [non sensu orig.,Syll. fung. 9:909 (1891)]. — Lectotypus: Winteria tuberculífera Ell. & Ev., i.e. N. grevillii.Coronophorella Höhn., Sitz.ber. Akad. Wiss. Wien, Math.-nat. Kl. 118(1): 1507 (1909). —Typus: Enchnoa chaetomioides Penz. & Sacc.Teratonema H. & P. Syd., Ann. Mycol. 15(3^1): 180 (1917). — Typus: Asterulacorniculariiformis P. Henn., i.e. N. acanthostroma.? Schizocapnodium Fairm., Proc. Rochester Acad. Sei. 6(3): 93 (1921). — Typus: Sch.sarcinellum Fairm. (comp, below p. 318).Calyculosphaeria Fitzp., Mycologia 15(2): 45 (1923). —Nov. nom. pro Winterella Berl.Biciliospora Petr., Sydowia 6(5-6):429 (1952). — Typus: B. velutina Petr. — Syn. ex descr.Scortechiniella Arx & Müll., Beitr. Krypt.-fl. Schweiz 11(1): 382 (1954). — Typus: Trichosphaerellasimilis Bres.Scortechiniellopsis Sivanesan, Trans. Brit. Mycol. Soc. 62(1): 36 (1974). — Typus: Sc.leonensis Sivanesan.Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 307An alphabetical list of the 22 accepted species follows, with nomenclators,published exsiccata (if any) and sketches of their distributions as well as short noteson their diagnostic features. Most of them are readily known by some unique featureor by a combination of two conspicuous features. Thus N. acanthostroma by the"spiny" subiculum branches, and N. callista by the bright (brown) colour of thesubiculum; N. parasitons by the small ascocarps (200), regularlyarranged spores; N. velutina and N. similis by the appendiculated spores, 8 and ca.32 per ascus respectively; N. tetraspora and N. collapsa by the fusiform, distinctly1-septated and eventually coloured spores, long and narrow and only 4 per ascusreaching maturity in the former, shorter and thicker in the latter; N. macrospora andN. gigantospora by the very large spores, 24^15x6-6.5 /xm and 36-45x12-14 /amrespectively.(1) Nitschkia acanthostroma (Mont.) Nannf., Sv. Bot. Tidskr. 69(1): 58 (1975).Sphaeria acanthostroma Mont., Ann. Sci. Nat. Bot. 4(3): 126 (1855); Syll. crypt, p. 226(1856). — Trichosphaeria acanthostroma Sacc., Syll. fung. 1:454 (1882). — Scortechiniaacanthostroma Sacc. & Beri., Atti R. 1st. Venet. Sci. Lett. Art. 6(3:4):714 (1885). — Byssosphaeria(Trichosphaeria) acanthostroma Cke, Grevillea 15(=n. 76): 122 (1887). —Tympanopsis acanthostroma Miill. & Arx, Phytopath. Zeitschr. 24(4): 365 (1955). — Typus:French Guiana, Leprieur 1163 (PC).Sphaeria culcitella Berk. & Rav. in Rav., F. carol. 4:53 (1855; nom. nud.); in Berk. & Curt.,Proc. Amer. Acad. Arts Sci. (Boston) 4:128 (1860). —Scortechinia culcitella Speg., Anal. Soc.Cie. Argentina 26:29 (1888). — Lectotypus: Rav., F. carol. 4:53.[Sphaeria aculeata Berk. & Br. ex Cke, Grev. 15(=n. 76): 122 (1887; nom. nud., ut syn.Byssosph. acanthostr.).]Astemia corniculariiformis P. Henn., Hedwigia 36(4): 218 (1897). — Teratonemacorniculariiforme H. & P. Syd., Ann. Mycol. 15(3-4): 180 (1917). — Typus:Brazil, S ,aCatarina, Moller 228 (S !).Orbicula Richenii Rick, Ann. Mycol. 2(3): 245 (1904). — Typus: Rick, F. austro-amer. 1.EXS.: EU. & Ev., N. Amer. F. 2:2356 (Trichosph. ac.) PAD!, UPS! — Rav., F. carol. 4:53(Sph. culc.) UPS!. — Rick, F. austro-amer. 1 (Orb. Rich.) PAD!, S!.[Non: Theiss., Dec. f. bras. 90 (Trichosph. ac.)=Chaetosphaerella sp.~\Subiculum forming large and dense blackish patches, the hyphal terminationscharacteristically spiny. Ascocarps often almost buried, 0.25-0.4 mm across. Asci8-sporous. Spores ellipsoid to ovoid, 6-8x3^1 p.m, hyaline (or eventually withpale-smoky walls?).DISTRIBUTION: Widespread in subtropical and tropical regions; known fromsouth-eastern North America (U.S.A.: Florida, Louisiana, South Carolina), Centraland South America (Nicaragua, French Guiana, Bolivia, Brazil, Paraguay), Africa(Sao Tomé, Sierra Leone, Uganda, Zaïre), Asia (Ceylon, Java, The Philippines)and Australia (Queensland).SvenskBot. Tidskr. 69 (1975)

308 J. A. Nannfeldt(2) Nitschkia affinis (H. & P. Syd.) Nannf. nov. comb.Fracchiaea affinis H. & P. Syd., Ann. Mus. Congo Bot. 5:16 (1909). — Typus: Zaire,Vanderyst (S!).Ascocarps 0.5-0.7 mm. Asci ca. 32-sporous. Spores subcylindrical to narrowlyellipsoid, straight to slightly curved, 5.5-7x 1-1.5 /xm, hyaline, with 2 drops.DISTRIBUTION: Tropical Africa (Zaire). Only the type collection.(3) Nitschkia brevispina (Münk) Nannf. nov. comb.Trichosphaeria brevispina Münk, Dansk Bot. Ark. 17(1): 185 (1957). — Typus: Denmark,Müller, Münk & Nannfeldt (C irrepertus 1973).Similar to N. grevillii.Ascocarps 0.3-0.4 mm; part of the peridial cells prolongedinto pointed spines (up to 20 /u,m long). Asci 8-sporous. Spores subcylindrical tonarrowly ellipsoid, slightly curved, (7-)10-16x3^t /u.m, hyaline, with 2 large drops,sometimes almost filling the lumen, eventually with a (pseudo?)septum.rare.DISTRIBUTION: Europe (Sweden, Denmark, Finland, England). Evidently very(4) Nitschkia broomeiana (Berk.) Nannf., Sv. Bot. Tidskr. 69(1): 60 (1975).Sphaeria (Caespitosae) broomeiana Berk., Hooker's Journ. Bot. 6:231 (1854). — Coronophorabroomeiana Sacc., Syll. fung. 1: 106 (1882). —Fracchiaea broomeiana Petch, Ann. R.Bot. Gard. Peradeniya 6(4): 333 (1917). — Typus: Ceylon, Thwaites (K fide Petch).? Sphaeria rasa Berk, in Hook., Fl. N. Zeal. 2:205 (1855). —Fracchiaea rasa Sacc., Syll.fung. 1:95 (1882). — Typus: New Zealand, Colenso (holotypus: K fide Fitzp.; isotypus: PAD!inutilis 1974).[Sphaeria subcongregata Berk. & Curt, in Rav., F. carol. 4:57 (1855; nom. nud.); ex Cke,Grevillea 15(=n. 75): 83 (1887; nom. nud., ut syn. Fr. subconnatae). —] Fracchiaea subcongregataEll. & Ev., N. Amer. Pyrenom. p. 244 (1892). — Typus: Rav., F. carol. 4:57.Fracchiaea heterogenea Sacc., Atti Soc. Ven.-Trent. Sei. Nat. 2(2): 163 (1873). — Coll.orig.: "in agro Tarvisino et Patavino, vere, frequens".Echusias Vitis "(Schulzer)" Hazsl., Verh. zool.-bot. Ges. Wien 23:367 (1873) [non GibberaVitis Schulzer (1870)]. — Fracchiaea Vitis Höhn., Ann. Mycol. 17(2-6): 131 (1920). — Typus:Czechoslovakia, Schulzer v. Müggenburg (n.v.). — Syn. fide Höhnel I.e.[,Sphaeria brevibarbata Berk. & Curt, in sched. —] Cucurbitaria brevibarbata Berk. & Curt,ex Berk., Grevillea 4(=n. 30):47 (1875). —Fracchiaea(?) brevibarbata Sacc., Syll. fung. 1:94(1882). — Coll. orig.: USA, South Carolina, Ravenel 1558 & 1803 (K fide Fitzp.).Sphaeria subconnata Berk. & Curt, ex Berk., Grevillea 4(=n. 32): 141 (1876; nom. illegit.;non Sph. subconnata Schw., 1832). — Coelosphaeria subconnata Sacc., Syll. fung. 1:93(1882). —Fracchiaea subconnata Cke, Grevillea 15(=n. 75):83 (1887);Berl., Icon. fung. 3:24(1900). — Nitschkia subconnata O. Kze, Rev. gen. pi. 3(2): 501 (1898; "Nitschkea"). —Typus: USA, South Carolina, Ravenel (Curtis 2737 K fide Fitzp.).Gibbera moricarpa Cke, Grevillea 7(=n. 42): 51 (1878). —Fracchiaea moricarpa Sacc., Syll.fung. 1:94 (1882). — Coll. orig.: USA, Georgia, Ravenel 2471 & 2540 (K, NY fide Fitzp.).Fracchiaea cucurbitarioides Speg., Anal. Soc. Cie. Argentina 10(1): 16 (1880). — Typus:Argentina, Spegazzini (LPS fide Fitzp.).Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 309[,Sphaeria subconvexa Berk. & Rav. ex Cke, Grevillea 15(=n. 75):83 (1887; nom. nud., utsyn. Fr. moric.).]Fracchiaea americana Berl., F. moricolae 5:1 (1888; n.v.). — Typus: USA, New Jersey,Ellis 137 (isotypus: NY fide Fitzp.).Fracchiaea glomerata Pat., Journ. de Bot. 3(10): 168 (1889). — Typus: Venezuela, Gaillard285 (holotypus: PC fide Fitzp.; isotypi duo: S!).[Sphaeria pauridia Berk. & Curt, in sched. —] Nitschkia pauridia Cke, Grevillea 20(=n.96): 107 (1892). — Fracchiaea pauridia Berl., Icon. fung. 3:25 (1900). — Typus: USA, SouthCarolina, Curtis 1413 (isotypus: FH fide Fitzp.).Fracchiaea eucalyptina Berl., Icon. fung. 3:27 (1900). — Typus: Speg., Dec. myc. arg. 41.? Fracchiaea depressa Petch, Ann. R. Bot. Gard. Peradeniya 6(3): 221 (1917). — Typus:Ceylon, PDA 2922 (n.v.). — Syn. dub. fide Nannfeldt 1975:62.EXS.: Cav., F. longob. 119 (Fr. het.) BPI, FH, NY fide Fitzp.; S!, W! — Ell., N. Amer. F.692 (Gibb. moric.) CUP, NY fide Fitzp.; UPS! — Rav., F. amer. 343 (G. moric.) BPI, CUP,FH, NY fide Fitzp.; F. carol. 4:57 (Sph. subcongr.) BPI, CUP, FH, K, NY fide Fitzp.; UPS!— ? Roumeg., F. gall. 5568 (Fr. het.) UPS! inutilis. — Sacc., Myc. ven. 88 (Fr. het.) FH fideFitzp.; S!, W! — Speg., Dec. myc. arg 41 (Fr. het.) BPI, FH, NY fide Fitzp.; IMI!, PAD!,W!.[Non: Cav., F. Iongob. 234 (Fr. het.), q.e. Ditopella ditopa (Fr.) Schroet.]Ascocarps large (0.35-0.65 mm), only rarely collapsing into cupshape, mostly insmall pulvinate groups. Peridium with pointed spines (up to 25 pm long), sometimesforked and often with secondary "barbs". Asci large (up to 125x25 /am). Spores>200 per ascus, characteristically arranged, subcylindrical, slightly curved,8-11x1.5-2 /Ltm, hyaline, with one droplet in each end, eventually with a faintseptum.DISTRIBUTION: Widespread in most warmer countries. Known from Europe(France, Italy, Czechoslovakia, Yugoslavia. Records from more northern areas areerroneous, see Chapt. 8), North America (south-eastern USA, seen by me fromFlorida, Georgia, New Jersey, South Carolina, Virginia, recorded also fromLouisiana and Ohio, Fitzpatrick 1924: 106-107, and from North Carolina, Mycologia33(5): 572, 1941), Central and South America (Nicaragua, Venezuela, Brazil, Argentina),Africa (Sierra Leone, Ghana, Gambia, Southern Rhodesia, cp. Hopkins1938, 1939 and Doidge 1950:194), Asia (seen by me from India, Pakistan, Japan, andCeylon; China fide Teng 1934:368), Australia (South Australia, IMI!) and NewZealand (North Island).(5) Nitschkia callista (Berk. & Curt.) Nannf. nov. comb.[Sphaeria callista Berk. & Curt, in Rav., F. carol. 5:67 (1860; nom. nud. —] Cucurbitariacallista Eerk. & Curt, ex Berk., Grevillea 4(=n. 30):47 (1875). —Fracchiaea callista Sacc.,Syll. fung. 1:94 (1882). — Typus: USA, South Carolina, Curtis 342 (K).EXS.: Ell. & Ev., N. Amer. F. 1188 (Cucurb. c.) UPS!, W!; 2512 (Fracch. c.) PAD!, UPS! —Rav., F. carol. 5:67 (Sph. c.) UPS! — Rel. Farl. 26 (Fr. c.) S!, UPS!; 706 (Fr. c.) S!, UPS!, W!20-753873 Svensk Bot. Tidskr. 69 (19

310 J. A. NannfeldtAscocarps deeply cupulate, 0.25-0.4 mm, densely gregarious, surrounded by arather sparse, distinctly brown (not blackish) subiculum. Asci ca. 32-sporous.Spores suballantoid, slightly curved, 7-9x1.5 /im, hyaline with one drop in eachend.DISTRIBUTION: Seems confined to North America; seen by me only from theEast: Canada (Ontario) and USA (Alabama, Connecticut, Maryland, Pennsylvania,South Carolina, Virginia) but recorded also from the West (USA: Oregon, Zeller1927:130). — A record from Sweden is erroneous (see Chapt. 8). Judged from thedescription and illustration, a record (Teng 1936:506) from China (Hunan) is notconvincing. It may be an undescribed Species.(6) Nitschkia calyculus (Mont.) O. Kze, Rev. gen. pi. 3(2): 501 (1898; "Nitschkea")-,Berl., Icon. fung. 3:22 (1900).Sphaeria (Caespitosa) calyculus Mont., Ann. Sei. Nat. Bot. 2(14): 322 (1840); Syll. crypt, p.226 (1856). —Byssosphaeria (Caelosphaeria) calyculus Cke, Grevillea 15(=n. 76): 122 (1887).— Coelosphaeria (?) calyculus Sacc., Syll. fung. 9:444 (1891). — Winterina calyculus Höhn.Ann. Mycol. 16(1-2): 105 (1918). — Calyculosphaeria calyculus fitzp., Mycologia 15(2):51(1923). — Typus: French Guiana, Leprieur 372 (PC!).Similar to N. grevillii. Ascocarps 0.35-0.5 mm. Asci 8-sporous. Spores ellipsoid tosubcylindrical, almost straight, only 5-7x1.5-2 ¡im, the smallest amongst the8-sporous species, hyaline, with one droplet in each end.DISTRIBUTION: North America (USA: Florida, Petrak 1952c:400, as Cal.trist.; W!), Tropical South America (French Guiana) and Tropical Africa (Ghana,IMI!; Hughes 1953:11 as Cal.trist.).(7) Nitschkia chaetomioides (Penz. & Sacc.) Nannf. nov. comb.Enchnoa chaetomioides Penz. & Sacc., Malpighia 11(4): 390 (1897); Icon. fung. javan. p. 4(1904). —Coronophorella chaetomioides Höhn., Sitz. ber. Akad. Wiss. Wien, Math.-nat. Kl.118(1): 1507 (1909). — Scortechinia chaetomioides Arx & Müll., Beitr. Krypt.-fl. Schweiz11(1):381 (1954). — Tympanopsis chaetomioides Arx & Müll., Phytopath. Zeitschr. 24(4):365(1955). — Typus: Java, Penzig 260bis (holotypus: PAD!; isotypus: W!).Similar to N. acanthostromabut the dichotomously branched endings of thesubicular hyphae not spiny. Ascocarps 0.3-0.4 mm. Asci 8-sporous. Spores suballantoidto subreniform, 6-8x2-3 /im, hyaline, with one large drop in each end.DISTRIBUTION: According to Arx & Müller (1954:381) "Indischer Archipel,Afrika, Südamerika", often mistaken forN. acanthostromaand probably not rare inthe Tropics. Seen by me only from Africa (Ghana, Sierra Leone, IMI!) and Asia(Java).(8) Nitschkia collapsa (Rom.) Chen., Bull. Soc. Mycol. Fr. 34(1-2):73 (1918;"Nitschkea").SvenskBot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 311Bertia collapsa Rom., Bot. Not. 1889(1):24 (1889). — Herpotrichia collapsa Rehm,Hedwigia 42, Beibl. 4: (176) (1903). —Calyculosphaeria collapsa Fitzp., Mycologia 15(2): 52(1923). — Typus: Rom., F. exs. scand. 70.Sphaeria tristis Tode var. sporidiis majoribus Berk. &Br., Ann. Mag. Nat. Hist. 4(7): 435(1871). _ Coll. orig.: Hainault Forest ii. 1859 & Batheaston 22.iii. 1869 (vide Chapt. 8).Trichosphaeria vagans Boud., Icon, mycol. 4:339 (1910). — Typus: France, Boudier (PC).EXS.: Rom., F. exs. scand. 70 (B. coll.) FH fide Fitzp.; S!, UPS! — Roumeg., F. gall. 3524(iCoel. cup. ) COI!, UPS![Non: Weese, Eumyc. sel. 418 (Calyc. coll.) = Ac. tristis.]Similar to N. grevilliibut ascocarps larger, 0.45-0.75 mm. Asci 8-sporous, with adistinct apical wall-thickening and a subapical cylinder. Spores fusiform, straight,12-18x3.5-7 /im, initially with 4 large drops and hyaline wall, eventually with astrong median septum and smoky-grey wall.DISTRIBUTION: Seen by me from Europe (Sweden, Denmark, England,Belgium, France) and Australia (S. Australia). Recorded also from New Zealand(North Island, Fitzpatrick 1923:53). — The Australian specimen (G. Beaton 39, IMI116 884!) deviates by slightly longer spores (16-23x6-6.5 /xm) but is otherwisetypical.(9) Nitschkia confertula (Schw.) Nannf., Sv. Bot. Tidskr. 69(1): 59 (1975).Sphaeria confería Schw., Sehr. Nat. Ges. Leipzig 1:45 (1822); Fr., Syst. myc. 2:444 (1823)[non Sph. confería Fr., I.e. p. 435, q.e. Pyrenobotrys confería (Fr.) Theiss. & H. Syd.; videNannfeldt 1975:59]. —Byssosphaeria (Amphisphaeria) confería Cke, Grevillea 15(=n. 75): 81(1887; non sensu Cke). —Amphisphaeria confería Sacc., Syll. fung. 9:747 (1891; non sensuSacc.). — Typus: USA, North Carolina, Schweinitz (holotypus: PH; isotypi: BPI, FH fideFitzpatrick 1923:55-57 et Shear 1939:324; UPS!).Sphaeria conferiula Schw., Trans. Amer. Phil. Soc. 2(4): 211 (1832). — Tremalosphaeriaconferlula Ell., Proc. Acad. Nat. Sei. Philadelphia 1895:25 (1895; non sensu Ell., q.e.Chaeíosphaerella fusca). —Nom. nov. pro Sphaeria confería Schw. (non Fr.).Sphaeria euomphala Berk. & Curt, in Rav., F. carol. 4:54 (1855: nom. nud.); ex Berk.,Grevillea4(=n. 32): 141 (1876). — Boiryosphaeria euomphala Sacc., Syll. fung. 1:462 (1882).— Byssosphaeria (Eu-Rosellinia) euomphala Cke, Grevillea 15(=n. 76): 122 (1887). —Nilschkia euomphala Ell. & Ev., N. Amer. Pyrenom. p. 246 (1892). — Tympanopsiseuomphala Starb., Bih. K. Sv. Vet.-Akad. Handl. 19(3:2):24 (1894). — Scorlechiniaeuomphala Arx & Müll., Beitr. Krypt.-fl. Schweiz 11(1):380 (1954). — Coll. orig.: Rav., F.carol. 4:54.[.Sphaeria infroflexa Berk. & Rav. in sched. — Fracchiaea iniroflexa Cke, Grevillea 15(=n.75):84 (1887; nom. nud.). — Coll. orig.: Ravenel 1307, FH fide Fitzp.][Sphaeria craierella Berk. & Rav. ex Cke, Grevillea 15(=n. 76): 122 (1887; nom. nud., utsyn. Byssosphaeriae euomphalae). — Coll. orig.: Ravenel 1307; FH fide Fitzp.]Coelosphaeria beccariana Berl. & Pegl., N. Giorn. Bot. Ital. 24 (3): 110 (1892). —Nilschkiabeccariana O. Kze, Rev. gen. pi. 3(2): 501 (1898: "Nilschkea") — Typus: Italy, Beccari (isotypus:PAD! ut Coel. pisana, comp. Nannfeldt 1975:62).EXS.: Rav., F. carol. 4:54 (Sph. euomph.) FH, MO, NY, PH fide Fitzp.; UPS!SvenskBot. Tidskr. 69 (1975)

312 J. A. NannfeldtSimilar to N. grevillii. Ascocarps 0.3-0.5 mm. Asci 8-sporous. Spores ovoid toellipsoidal to subreniform, mostly inequilateral, 7-11 X3.5-5 /xm, with 2 large, oftenconfluent drops; wall early smoky-grey. — Mostly associated with Hypoxylonrubiginosum (Pers. ex Fr.) Fr. s. lat.DISTRIBUTION: Europe (Great Britain, Italy), North America (seen by me fromUSA: Georgia, Maryland, New York, North Carolina, South Carolina) and Africa:(Ghana, IMI!; South Africa fide Miller 1942:259 et Doidge 1950:146). Recorded alsofrom Central America (Arx & Müller 1954:380) and Asia (China: Chekiang andYunnan, Teng 1934:370).(10) Nitschkia cupularis (Fr. ex Fr.) Karst., Mycol. fenn. 2:81 (1873) [non sensuKarst., q.e. N.parasitans].Sphaeria cupularis Fr. [Sv. Vet.-Acad. Handl. 37:112 (1817)] ex Fr., Syst. myc. 2:416 (1823)[vix Pers., vide Nannfeldt 1975:53; nec Sphaeria (Poronia) cupularis Fr. 1830]. —Cucurbitaria cupularis S. F. Gray, Nat. Arr. Brit. PI. 1:519 (1821); Cke, Handb. Brit. F. p. 842(1871). — Cyathisphaera cupularis Dumort., Comment, bot. p. 87 (1822). — Hypoxyloncupulare Kickx, Fl. crypt. Louvain p. 114 (1835; n.v.). —Coelosphaeria cupularis Karst.,Medd. Soc. F. FI. Fenn. 5:42 (1879). — Tympanis cupularis Wallr., Fl. crypt. Germ. 2:428(1833; non sensu Wallr.). — Typus (sei. Fitzp. 1923, comp. Nannfeldt 1975:55): Fr., Sei. Suec.(ed. 1)231 (hololectotypus: FH; isolectotypus: UPS!).Melanopsamma numerosa Fautr. inRoumeg., Rev. Myc. 13(=n. 50): 76 (=Roumeg., F. exs.5629) (1891). — Typus: France, Fautrey iv: 159 (isotypi duo: UPS!; Roumeg., F. exs. 5629).Herpotrichia rehmiana P. Henn. & Kirschst. in P. Henn., Verh. Bot. Ver. Brandenb.40: xxviii (1898). — Typus: Germany, Kirschstein (isotypi duo: S!).EXS.: Fr., Scler. Suec. (ed. 1) 231 (Sph. cup.) FH fide Fitzp.; UPS! — Plowr., Sph. brit.1:63 (Sph. trist.) FH fide Fitzp.; S! — Rehm, Ascom. 1743 (N. trist.) BPI, FH fide Fitzp.; S!,W! — Roumeg., F. gall. 1551 (Chaetosph. trist.) C! inutilis, UPS!; F. exs. 5629 (Mel. num.)UPS![Non: Cke, F. brit. 2:561 (Cucurb. cup.)=N. par. — Desm., PI. crypt. 2:780(Sph. cup.)=N.par. — Fr., Scl. Suec. (ed. 2) 231 (Sph. cup.)=N. par. — Fuck., F. rhen. 968(Cue. cup.)=Wo.Barb.-Boiss. 591 (Coel. cup.)=N. par. — Holl & Schm., Deutschl. Schw. 6(Sph. cup.)=Cuc.(laburni?) + Camarosporium (comp. Nannfeldt 1975:54). — Karst., F. fenn. 861 (Sph.cup.)=N. par. — Kochm., Myc. pol 326 (N. cup.)=N. par. — Moug. & Nestl., St. crypt,vog.-rhen. 771 (Sph. cup.)=N. par. — Petr., Fl. boh.-mor. 2:1: 1634 (A'. cup.)=N. par.; Myc.gen. 177(N. cup.)=N. par. — Plowr., Sph. brit. 1:57 (Cue. cup.)=N. par. — Rehm, Ascom.983 (N. cup.)=N. par. — Roumeg., F. gall. 3524 (Coel. cup.)=N. coll. — Syd., Myc. march.1915 (N. cup.)=N. par.; 2430 (N. cup.)=N. par. — Vize, Micro-f. brit. 161 (Cue. cup.)=N.par. — Westend., Herb, crypt. 528 (Hyp. cup.)=N. grev.]Similar to N. grevillii.Ascocarps 0.3-0.45 mm. Asci 8-sporous. Spores subcylindrical,slightly curved to almost straight, 10-18x2-2.5 (im, hyaline, with 4 largedrops.DISTRIBUTION: Due to the bad confusion about this name only records substantiatedby specimens seen by Fitzpatrick or me can be taken into account. TheSvensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 313known area becomes thus restricted to Europe (Sweden, Great Britain, France,Germany, Austria).(11) Nitschkia floridana Fitzp., Mycologia 15(1): 31 (1923).Typus: USA, Florida, Thaxter (FH!).Similar to N. grevillii, but ascocarps more coarsely rough, 0.35-0.6 mm. Asci8-sporous. Spores suballantoid, 15-20x3^1.5 /um, hyaline with 2-4 drops, of whichthe two central may be very large, eventually with a distinct septum.DISTRIBUTION: North America (USA: Florida) and South America (Brazil: RioGrande del Sul 1904 Rick 7 as Bertia submoriformis, S!). Reported also from Asia(China: Yunnan, Teng 1934:368). — The type of S ph. submoriformis Plowright(1876:74) from California has not been seen by me. Should it turn out to beconspecific with the, present species, it would afford the correct epithet for it andwiden its known distribution considerably.(12) Nitschkia gigantospora Nannf. nov. nom.Calyculosphaeria macrospora Fitzp., Mycologia 15(2): 53 (1923) [non Nitschkia macrosporaTeng, 1934], — Typus: USA, Florida, Thaxter (FH!).Subiculum scanty. Ascocarps 0.5-0.6 mm. Asci 8-sporous. Spores cylindrical withrounded ends, straight, extremely large (36^15x12-14 pm), hyaline, with 2 verylarge drops, at maturity often less conspicuous, eventually relatively thick-walledand with a distinct septum.DISTRIBUTION: North America (USA: Florida). Only the type collection.(13) Nitschkia grevillii (Rehm in Starb.) Nannf., Sv. Bot. Tidskr. 69(1): 53 (1975).Melanopsamma Grevillii Rehm in Starb., Bih. K. Sv. Vet.-Akad. Handl. 16(3:3): 5 (1890). —Typus: Sweden, Grevillius (S!)."Nitschkia tristis (Pers.) Fuck.", Symb. mycol. p. 165 (1870), sensu Fuck., Wint. et auctt.plur. [non sensu orig., q.e. Acanthonitschkea tristis]. — Calyculosphaeria tristis ("Fuck.")Fitzp., Mycologia 15(2): 48 (1923). — Typus: Rbh., F. eur. 632 (comp. Nannfeldt 1975:57).Winteria tuberculífera EU. & Ev., Proc. Acad. Nat. Sei. Philadelphia 1890:240 (1890). —Winterella tuberculífera Berl., Icon. fung. 1:94 (1892; "tuberculigera"). — Winterinatuberculífera Sacc., Syll. fung. 14:589 (1899). — Typus: Canada, Dearness 1553 (holotypus:NY; isotypus: FH fide Fitzp.). — Syn. fide Fitzp.Wallrothiella fraxinicola Feltg., Ree. Mém. Trav. Soc. Bot. Luxembourg 16:286 (1905). —Typus: Luxembourg, Noppeney (LUX, comp. Nannfeldt 1975:62-63). — Syn. fide Höhnel1906:1215.Nitschkia winteriana Sacc., Atti Mem. R. Accad. Sei. Lett. Art. Padova 33:159 (1917; n.v.).— Typus: France, Flageolet (PAD!).EXS.: Fuck., F. rhen. 947 p.p. (Sph. trist.) S! (una cum Chaetosphaerella phaeostr.) [FHChaet. phaeostr. solum fide Fitzp.]=Hb. Barb.-Boiss. 590 p.p. (Coel. trist.) FH fíde Fitzp.; S!,20-753873 Svensk Bot. Tidskr. 69 (1975)

314 J. A. NannfeldtUPS! p.p. (una cum Chaet. phaeostr. et Ac. trist.), W! — Rbh., F. eur. 632 (Sph. trist). CUP,FH, NY fide Fitzp.; S!, UPS!, W! — Vize, Micro-f. brit. 391 FH fide Fitzp. — Westend., Herb,crypt. 528 (Hypox. cup.) COI!, LD! 1Subiculum scanty to profusely developed but never hiding the ascocarps. Thesemedium-sized (0.3-0.5 mm), gregarious to scattered. Asci 8-sporous. Spores subcylindricalto subfusoid, mostly straight, 6-9x1.5-2.5 ¡j,m, hyaline, with 2(-4) largedrops, eventually with a (pseudo?)septum.DISTRIBUTION: This seems to be one of the least rare and most widespreadtemperate species of the genus, being known with certainty from Europe (Sweden,Denmark, Norway, England, Belgium, Luxembourg, France, Germany, Switzerland),North America (Canada: Ontario and USA: Florida, Michigan, New York),and South America (Trinidad, Fitzpatrick 1923:51). It is also recorded from Poland(Schroeter 1894:313) and Italy (Traverso 1907:359) as N. tristis and from Asia(China: Chekiang and Yunnan, Teng 1934: 369) as Cal. tristis. — The record of Cal.tristis from Africa (Ghana) by Hughes (1953:11) refers to N. calycuius(q.v.)(14) Nitschkia leonensis (Sivan.) Nannf. nov. comb.Scortechiniellopsis leonensis Sivan., Trans. Brit. Mycol. Soc. 62(1):37 (1974). — Typus:Sierra Leone, Deighton M 6233 (IMI!).Similar to N. acanthostroma but the repeatedly dichotomously branched subicu-Iar hyphae with pale, acuminate (not spiny) endings. Ascocarps 0.3-0.4 mm. Asci ca.32-sporous. Spores broadly allantoid to subreniform, 5-7x2-3 /xm, hyaline, with 2large, often confluent drops, sometimes with a (pseudo?)septum.DISTRIBUTION: Tropical Africa (Sierra Leone). Only the type locality (3 finds,all on Cynometra leonensis).(15) Nitschkia macrospora Teng, Sinensia 4(12): 368 (1934).Typus: Yunnan, Tsiang 301 (n.v.).No subiculum. Ascocarps "seated on a pseudoparenchymatous stroma",0.35-0.55 mm, coarsely roughened, collapsing. Asci 8-sporous. Spores allantoid,occasionally straight, very large (26-45x6-6.5 /xm), hyaline, guttulate, eventuallywith a pseudoseptum. (Compiled from the diagnosis.)DISTRIBUTION: Asia (China: Yunnan). Only the type collection.(16) Nitschkia multiasca (Pat. & Gaill.) Nannf. nov. comb.Fracchiaea(?) multiasca Pat. & Gaill., Bull. Soc. Mycol. Fr. 4(3): 106 (1889). — Typus:Venezuela, Gaillard 1:24 p.p. (n.v.; comp. Fitzpatrick 1924:109).1 It seems to have remained unobserved that there exist two variants (editions?) of this exsiccatum. Thebound copy in LD has on the printed label a more detailed locality than has the loose copy in COI.SvenskBot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 315Ascocarps 0.25 mm, collapsing into cupshape, surrounded by brown hyphae. Ascica. 32-sporous. Spores ovoid, straight, 5-6x3 /am, hyaline, with 2 droplets. (Compiledfrom the diagnosis.)DISTRIBUTION: Tropical South America (Venezuela). Only the type collection.(17) Nitschkia parasitans (Schw.) Nannf., Sv. Bot. Tidskr. 69(1): 52 (1975).Sphaeria parasitans Schw., Trans. Amer. Phil. Soc. 2(4): 206 (1832). — Typus: USA:Pennsylvania, Schweinitz (holotypus: PH inutilis fide Ell. & Ev. 1892:794; isotypus: UPS!).Nectria (Gibbera) Hippocastani Otth, Mitth. naturf. Ges. Bern 1868:57 (1869; nom. confus.vide Nannfeldt 1975:63). —Calonectria (Chiajaea)Hippocastani Sacc., Hedwigia 35, Beibl. p.xxxiii (1896). — Typus: Switzerland, Otth (n.v.). — Syn. fide Höhnel 1920.Nitschkia Fuckelii Nke in Fuck., Symb. Mycol. p. 165 (1870; non rite pubi.). —Coelosphaeria Fuckelii Sacc., Atti Soc. Ven. Trent. Sci. Nat. 2:163 (1873; non rite pubi.).Typus: Fuck., F. rhen. 968 ("Cucurbitaria cupularis").Coelosphaeria anceps Sacc. & Malbr., Atti R. 1st. Veneto Sci. Lett. Art. 6(1:2); 1273 (1883).—Nitschkia anceps Lindau in Engl. & Prantl, Nat. Pfl.-Fam. 1(1): 409 (1897); Beri., Icon. fung.3:23 (1900). — Typus: France, Malbranche 512 (PAD!).Winteria tuberculifera E. & E. var. caespitosa E. & E., N. Amer. Pyrenom. p. 212 (1892).Typus: North America (NY), — Syn. fide Fitzp.Nitschkia moravica Niessl in Paul, Verh. Naturf. Ver. Brünn 47:138 (1909). — Typus:Czechoslovakia, Paul (n.v.). — Syn. ex descr.EXS.: Berk., Brit. f. 174 (Sph. acervata) FH, MO fide Fitzp.; UPS! — Cke, F. brit. 2:561(Cucurb. cup.) IMI!, UPS! —Desm., PI. crypt. 2:780 (Sph. cup.) UPS!, W! — Fr., Sci. Suec.(ed. 2) 231 (Sph. cup.) UPS! — Fuck., F. rhen. 968 (Cue. cup.) FH fide Fitzp.; S!;=Hb.Barb.-Boiss. 591 (Coel. cup.) FH fide Fitzp.; S!, UPS!, W! — Karst., F. fenn. 861 (Sph. cup.)FH fide Fitzp.; UPS! — Kochm., Myc. pol. 326 (N. cup.) UPS! — Moug. & Nestl., St. crypt,vog.-rhen. 771 (Sph. cup.) NY fide Fitzp.; S!, UPS!, W!; 1455 (Sph. cucurb.) UPS ! — Petr., Fl.boh.-mor. 2:1:1634 (N. cup.) C!, S!; Myc. gen. 177 (N. cup.) IMI!, S!, UPS!, W! — Plowr.,Sph. brit. 1:57 (Cue. cup.) S! —Rehm, Ascom. 983 (N. cup.;=Syd., Myc. march. 1915) H! S!,W! — Roumeg., F. gall. 1488 (N.fuck.) CUP, FH fide Fitzp.; COI!, UPS! — Sacc., Myc. ven.1449 (Coel. fuck.) UPS!, W! —Syd., Myc. march. 1915 (N. c«p.;=Rehm, Ascom. 983) FH, NYfide Fitzp. ; S!, UPS !, 2430 (N. cup.) FH, NY fide Fitzp. ; S !, UPS !; 4131 (N. trist.) FH, NY fideFitzp.; B!, S!, UPS! — Thüm., Myc. univ. 1947 (N. fuck.)BPl, FH, NY fide Fitzp.; B!, COI!,UPS!, W! — Vize, Micro-f. brit. 161 (Cue. cup.) IMI!Ascocarps small (0.2-0.3 mm), mostly caespitose on stromata of Nectriacinnabarina.Asci 8-sporous. Spores suballantoid to boomerang-shaped, 9-16x2-3 pm,hyaline, with one droplet (rarely 2) in each end, eventually with a faint(pseudo?)septum.DISTRIBUTION: Known with certainty from Europe (Sweden, Denmark, TheFaeroes, Finland, Norway, England, Belgium, France, Germany, Switzerland, Austria,Poland, Czechoslovakia , Italy, S.S.S.R.) and North America (USA: Pennsylvania).Recorded also from USA: Montana (Ellis & Everhart 1892:245) and fromAsia (India, several finds; Mundkur & Ahmad 1946:5). This species, rather con-SvenskBot. Tidskr. 69 (1975)

316 J. A. Nannfeldtspicuous in spite of the minuteness of its ascocarps, appears evidently more frequentlyin certain years (springs) and may then be fairly copious in its localities.(18) Nitschkia pezizoidea (Pat. & Gaill.) O. Kze, Rev. gen. pi. 3(2): 501 (1898;"Nitschkeapezizodea").Coelosphaeria pezizoidea Pat. & Gaill., Bull. Soc. Mycol. Fr. 4(3): 106 (1889;"Caelosphaeria"). — Winterina (?) pezizoidea Höhn., Ann. Mycol 16(1-2): 105 (1918). —Typus: Venezuela, Gaillard 266 (PC).Similar to N. grevillii. Ascocarps 0.35-0.45 mm, collapsing into deeply cupulate,apical papilla unusually prominent. Asci 8-sporous. Spores fusiform to ovoid,straight, 7-11x2-3 fxm, hyaline, eventually with a distinct septum. (Compiled fromFitzpatrick 1923:52.)DISTRIBUTION: South America (Venezuela). Only the type collection.(19) Nitschkia similis (Bres.) Nannf. nov. comb.Trichosphaerella similis Bres., Bull. Jard. Bot. Bruxelles 4(1);7 (1914). — Scortechiniellasimilis Arx & Müll., Beitr. Krypt.-fl. Schweiz 11(1): 383 (1953). — Typus: Zaïre, Vanderyst(S!).Similar to N. acanthostromabut the dichotomously branched endings of thesubicular hyphae not spiny. Ascocarps 0.3-0.5 mm. Asci ca. 32-sporous. Sporesreniform to suballantoid, often also spirally twisted (up to 90°), hyaline, with afiliform, ca. 5 /xm long curved appendix at each end, 6-9x3 /xm, and with 2 largedrops, almost filling the lumen, and eventually with a distinct (pseudo?)septum.DISTRIBUTION: Tropical Africa (Zaïre). Only the type collection.(20) Nitschkia tetraspora Nannf. nov. spec.N. collapsa similis et certe affinis. Differt praecipue sporis angustioribus et longioribus(15-22x2.5-3 /urn), initio octonis in asco sed normaliter quaternis tantum maturescentibus.Ascocarpia (sicca) cupulato-collapsa (0.4-0.5x0.25-0.3 mm), papilla apicali parva inconspicua.Sporae anguste fusiformes, rectae vel interdum leviter curvatae, 1-septatae, pariete deniquefuscescente.Typus: Kenya, Nannfeldt 22058 (holotypus: UPS; isotypus: S).Subiculum absent or scanty. Ascocarps scattered to densely gregarious, seated on peridermor on inner-bark beneath loosening parts of the periderm, turbinate and (when dry) collapsed tocupshape, 0.4-0.5x0.25-0.3 mm, the apical papilla very small. Asci 25-30x8-11 /x (p. sp.),early deliquesent, initially 8-spored, but as a rule only (the upper) 4 spores reach maturity.Spores fusiform, straight or sometimes slightly curved, 15-22x2.5-3 ¡xm, with initially 4-6drops, later confluent to 2 and at a certain stage almost filling the lumen, successively getting amedian septum and constriction; wall smooth, eventually smoky-grey. Abortive spores acicularto narrowly fusiform (ca. 16x 1-1.5 /xm), hyaline, 1-septate.DISTRIBUTION: Tropical Africa (Ghana, Kenya, Sierra Leone).SvenskBot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 317Fig. 4. Nitschkia tetraspora (holotypus). fa) Substrate with dry ascocarps (ca. 8x; phot. K. Holm). — (b, c)Spores in the stage they become free; in the upper right comer of b four abortive spores are seen surroundinga normal (ca. 900 x).Africa: Kenya: Coast Prov., Kwale Distr., Jadini Forest 26.ii.1972 J. A. Nannfeldt 22058.(Type.)Ghana: Amanase 7.vi.l949 S. J. Hughes 1172 (IMI 38190a!); Asuansi (Cola nitida) 9.vi.l949S. J. Hughes 1183 (IMI 44569!); Bunsu (Hevea bras.) 17.vi. 1949 S. J. Hughes 1331 (IMI38076c!).Sierra Leone: Puabu ("Kpuabu"; Gaura) (Theobroma cacao) 29.ix. 1960 A. B. KattaM. 6605 (IMI 84483!).(21) Nitschkia uniseriata (Fitzp.) Nannf. nov. comb.Tympanopsis uniseriata Fitzp., Mycologia 15(2):58 (1923). — Typus: USA: Florida, Thaxter(FH!).Subiculum dense. Ascocarps 0.25-0.35 mm, cupulate, gregarious to scattered.Asci 8-sporous, cylindrical (p. sp. 60-70x8-10 /xm). Spores uniseriate, often transverselyorientated, broadly ellipsoid to ovoid, slightly inaequilateral, 7-9x4.5-5.5¿¿m with 2 large often confluent drops, wall before maturity hyaline with distinctcyanophilous warts, eventually minutely echinulate, smoky- to yellowish-brown.DISTRIBUTION: North America (USA: Florida). Only the two finds listed byFitzpatrick.20-753873 Svensk Bot. Tidskr. 69 (1975)

318 J. A. Nannfeldt(22) Nitschkia velutina (Petr.) Nannf. nov. comb.Biciliospora velutina Petr., Sydowia 6(5-6):429 (1952). — Typus: Puerto Rico, Earle 284(n.v.).Similar to N. acanthostroma.Subicular hyphae dichotomously branched, theirterminations short, often conical or subglobose (not spiny). Ascocarps 0.5-0.6 mm.Asci 8-sporous. Spores narrowly ellipsoid to subcylindrical, mostly straight,19-30x4.5-7 ¡xm, hyaline, with numerous droplets and at each end with a filiform(ca. 10 ftm long) appendix, curved backwards. (Compiled from the diagnosis.)DISTRIBUTION: Tropical America (Puerto Rico). Only the type collection.The above list is not exhaustive. There hide certainly in all warmer countries andin the South Temperate Zone a number of undescribed species as well as suchdescribed in the most unexpected genera. Also several (non-European) fungi knownto me only from literature may be valid species, although their true affinities andtheir distinctness remain doubtful until their types have been examined, e.g. thefollowing:Fracchiaea australis Speg. in sched., illustrated by Fitzpatrick (1924: pi. 10 fig. 5) and due toits "Quellkorper" regarded by him (I.e. p. 108) as a Cryptosphaerella.Coelosphaeria corticata Ell. & Ev. from USA (Missouri), regarded by Fitzpatrick (1923:38)after studying type material as a Coronophorella (asci 8-sporous, spores "moderately curved,10-14x3 ft").Tympanopsis coelosphaerioides Penz. & Sacc. var. minor Teng (1932). The main species isthe bitunicate Auerswaldia examinans (Mont. & Berk.) Sacc. (cp. Chapt. 4), but according toTeng's description and illustration (1934:370) his variety seems to be widely different andperhaps a true Nitschkia, marked by deeply cupulate ascocarps (0.28-0.45 mm), 8-sporousasci, mostly uniseriate, ellipsoid to inaequilateral spores, 11-15x4.5-6.5 pm, usually uniguttate,with eventually smoky-yellow wall.Nitschkia hainanensis Teng & Ou in Ou (1936). The description and illustration of thisspecies from China (Hainan) indicate a Nitschkia, close to N. floridana but probably distinct,the main differences being the minute roughness of the ascocarps and the larger spores(18-32x4-5 ixm).Schizocapnodium sarcinellum Fairm. (1921). The very unsatisfactory type specimen (fromUSA: New York) was studied by Petrak (1952a:265). From his description and drawingsMiiller & Arx (1962:819) suggested that it might be congeneric with Gaillardiella pezizoidea(cp. Chapt. 4 above). If it is Coronophoralean at all, it is in my eyes rather a species ofNitschkia, as there is no sign of the ringshaped thickening of the peridium, characteristic ofGaillardiella. The "double-spores" are clearly an artifact due to beginning decay and the verydark spore wall may have the same cause. An exacter disposition may perhaps be possiblewhen the mycoflora of the type area has become better known.6. The Species of AcanthonitschkeaThe changed delimitation of Acanthonitschkea, as motivated in Chapt. 4, with thepresence (at least on the subiculum) of strong, dark, unseptate bristles as the leadingSvensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 319character results in the following nomenclátor and in the subsequent alphabetical listof the accepted four species with their nomenclators, exsiccata, diagnostic featuresand distributions, all in the same manner as for the Nitschkiae in the precedingchapter.Acanthonitschkea Speg., Anal. Mus. Nac. Bueno Aires 3(10): 116 (1908). — Typus:Ac. argentinensisSpeg.Euacanthe Theiss., Ann. Mycol. 15(3-4): 272 (1917). — Typus: Meliolopsis usambarensisRehm, i.e. Ac.foveolata.Neotrotteria Sacc., Bull. Orto Bot. R. Univ. Napoli 6:45 (1918). — Typus: N. pulchellaSacc.Petelotia Pat., Bull. Soc. Mycol. France 40(1): 35 (1921). — Typus: P. tonkinensis Pat., i.e.Ac. pulchella.Sydowinula Petr., Ann. Mycol. 21(3-4):277 (1923). — Typus: S. moravica Petr., i.e. Ac.tristis.? Fitzpatrickia Cif., Mycologia 20(1): 29 (1928). — Typus: F. massae Cif. (prob. =Ac.foveolata). — Syn. dub. ex descr. (comp. Nannfeldt 1975:60).The species are all well marked and readily known from each other. Three ±tropical species have bristles also on the ascocarps, one (Ac. pulchella)polysporous (ca. 32-sporous) asci, and the others 8-sporous (Ac. argentinensissuballantoid, strongly curved spores and Ac. foveolata with ± ellipsoidal and ±straight spores). The fourth, so far exclusively European species (Ac. tristis) has nobristles on the very ascocarps but a "crown" of such round their bases.haswithOnly one additional species has been described under any of the relevant genericnames, viz. Ac. coloradensis Cash & Davidson (1940), but as shown in Chapt. 3(Nannfeldt 1975:62) this is decidedly misplaced and rather of Diaporthalean affinity.(1) Acanthonitschkea argentinensis Speg., Anal. Mus. Nac. Buenos Aires 3(10): 116(1908).Typus: Argentina, Spegazzini 2107 (LPS!).Ascocarps 0.2-0.3 mm, with bristles 100-200 pm long. Asci 8-sporous. Sporessuballantoid, strongly curved, often almost semicircular or horseshoe-shaped,5-8x2 /u.m, hyaline, 4-guttate.DISTRIBUTION: America (Puerto Rico fide Fitzpatrick 1923:63, Argentina) andAfrica (Ghana, IMI!; Hughes 1953:4).(2) Acanthonitschkea foveolata (Berk. & Curt, ex Berk.) Nannf., Sv. Bot. Tidskr.69(1): 60 (1975).Sphaeria (Byssisedae) foveolata Berk. & Curt, ex Berk., J. Linn. Soc. 10:387 (1868). —Rosellinia foveolata Sacc., Syll. fung. 1:271 (1882). — Coll. orig.: Cuba, Wright 489 & 548(spec, auth.: LPS 5810!).20-753873 Svensk Bot. Tidskr. 69 (1975

320 J. A. NannfeldtSphaeria (Byssisedae) hystricula Berk. & Br., J. Linn. Soc. 14:125 (1873). —Roselliniahystricula Sacc., Syll. fung. 1:274 (1882). —Chaetosphaeria hystricula Cke, Grevillea 15(=n.76): 124 (1887). — Fracchiaea hystricula Petch, Ann. R. Bot. Gard. Peradeniya 5(4): 290 (1912;non sensu Petch, q.e. Ac. pulchella). — Typus: Ceylon, Thwaites 171 p.p. = 1074 1 (isotypus:UPS!).["Trichosphaeria acanthostroma (Mont.) Sacc." sensu Speg., Anal. Soc. Cie. Argentina26:28 (1888). — Cp. Nannfeldt 1975:58.]Meliolopsis usambarensis Rehm in P. Henn., Deutsch Ost-Afrika 5C:31 (1895). —Euacanthe usambarensis Theiss., Ann. Mycol. 15(3-4):272 (1917). — Scortechiniausambarensis Arx & Müll., Beitr. Krypt.-fl. Schweiz 11(1): 381 (1954). — Tympanopsisusambarensis Müll. & Arx, Phytopath. Zeitschr. 24(4):365 (1955). — Typus: Burundi, Hoist(S!).Acanthonitschkea macrobarbata Fitzp., Mycologia 15(2):63 (1925). — Typus: Trinidad,Thaxter (FH!).? Fitzpatrickia Massae Cif., Mycologia 20(1):30 (1928). — Scortechinia Massae Arx &Müll., Beitr. Krypt.-fl. Schweiz 11(1): 381 (1954). — Tympanopsis Massae Müll. & Arx,Phytopath. Zeitschr. 24(4): 366 (1955). — Typus: loc., temp, et matr. ignot., leg. C. Massa(School of Viticulture and Enology, Alba, Italy; n.v.). — Cp. Nannfeldt 1975:60.Ascocarps 0.25-0.5 mm with bristles 200-300 /tm long. Asci 8-sporous. Sporesovoid or ellipsoid to subfusiform, inequilateral, almost straight, 5-8x2-3 ¿¿m,hyaline (eventually with brownish wall?), with 2 drops, as a rule confluent to oneeccentrically placed.DISTRIBUTION: Known from the Americas (USA: South Carolina fide Berkeley1868:387, Cuba, Trinidad, Brazil, Paraguay), tropical Africa (Burundi), and tropicalAsia (Ceylon).(3) Acanthonitschkea pulchella (Sacc.) Nannf. nov. comb.Neotrotteria pulchella Sacc., Bull. Orto Bot. R. Univ. Napoli 6:45 (1918). — Typus:Singapore, Baker 5277 (n.v.; comp. Petrak 1962:353).["Fracchiaea hystricula (Berk. & Br.) Petch" sensu Petch, Ann. R. Bot. Gard. Peradeniya5(4):229 (1912); non s. orig., q.e. Ac.foveolata.]Petelotia tonkinensis Pat., Bull. Soc. Mycol. France 40(1):35 (1921). — Typus: NorthVietnam, Petelot 570 (holotypus: FH!).Ascocarps 0.3-0.5 mm with bristles 100-250 fim long. Asci ca. 32-sporous. Sporessuballantoid, 6-12x 1.5-2(-2.5) /u.m, hyaline, with 2(-3) droplets.DISTRIBUTION: Tropical Africa (Ghana, IMI!; Hughes 1953: 18 as Fr.hystr.)and tropical Asia (Ceylon, North Vietnam, Malaya). A sample (IMI 104 902) fromNorth Borneo may also belong here, but its stage is too advanced for certaindetermination.—The matrix is in most cases Heveabrasiliensis.1 Expressions such as "nos. 1074, 171 in part" frequently used by Berkeley & Broome in connection withThwaites' Ceylon fungi do not indicate two different collections but mean that part of a mixed collection (inthis case 171) was given a number (1074) of its own (Petch 1912:266).Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 321(4) Acanthonitschkea tristis (Pers. ex Fr.) Nannf., Sv. Bot. Tidskr. 69(1): 57 (1975).Sphaeria tristis Pers. [Icon, descr. fung. 2:49 (1800)] ex Fr., Syst. myc. 2:444 (1823) [nonSphaeria tristis Tode 1791, q.e. Chaetosphaerella phaeostroma fide Fuck.]. —Nitschkia tristisFuck., Symb. mycol. p. 165 (1870; non rite pubi, et non sensu Fuck., q.e. N. grevillii). —Coelosphaeria tristis Karst., Medd. Soc. F. Fl. Fenn. 5:42 (1879; non sensu Karst.); Sacc. inRoumeg. & Sacc., Rev. myc. 3(=n. 11):42 (1881; non sensu Sacc., q.e. N. cupularis s. orig.).— Byssonectria (Caelosphaeria) tristis Cke, Grevillea 15(=n. 76): 122 (1887; sensu Sacc.). —Winterina tristis Höhn., Ann. Mycol. 16(1-2): 105 (1918; non sensu Höhn.). — Lectotypus (sel.Nannfeldt 1975:57): Hb. Persoon L. 910, 269-833!Coelosphaeria media Sacc., Michelia2:592 (1882). —Nitschkia media O. Kze, Rev. gen. pl.3(2):501 (1898; "Nitschkea"). — Loranthomyces (?) media Höhn., Ann. Mycol. 16(1-2): 105(1918). _ Typus: Switzerland, Morthier 4 (PAD!). — Vide Nannfeldt 1975:61.Sydowinula moravica Petr., Ann. Mycol. 21(3-4):277 (1923). — Typus: Moravia, 18.vi. 1922,Petrak (n.v.). — Syn. ex descr. (comp. Nannfeldt 1975:63).EXS.: Cke, F. brit. 1:269 (Sph. tr.) UPS! — Fr., Sci. Suec. 386 (Sph. tr.) UPS! — Fuck., F.rhen. 947 p.p. (Sph. tr.) W! (copy with number and name in handwriting)=Hb. Barb.-Boiss. 590p.p. (Coel. tr.) UPS! (una cum N. grev. etChaet. phaeostr.). —Weese, Eumyc. sel. 448 (Calyc.coll.) UPS![Non: Plowr., Sph. brit. 1:63 (Sph. tr.)=N. cup. s. orig. — Rbh., F. eur. 632 (Sph. tr.)=N.grev. — Rehm, Ascom. 1743 (N. tr.)=N. cup. s. orig. — Roumeg., F. gall. 1485 (Sph. tr.)=l\1551 (Chaetosph. tr.)=N. cup. s. orig.]Bristles 150-200 /am long, only on subiculum but most thickset round the bases ofthe ascocarps, which are 0.3-0.35 mm. Asci 8-sporous. Spores subcylindrical tonarrowly ovoid, slightly curved, 6-9x 1.5-2 /am, hyaline, (2-)4-guttulate, eventuallywith an obscure (pseudo?)septum.DISTRIBUTION: Due to the confusion about this species, only specimens actuallystudied by me (plus of course Petrak's record of his new species) can be takeninto account, which gives an area restricted to Europe (Sweden, Norway, England,France, Germany, Switzerland, Czechoslovakia).7. Nitschkiopsis stictarum nov. gen. et sp.(with a contribution by R. Santesson)The observant reader may have missed any allusion in the preceding chapters to thenew African lichenicolous species. The reason is that the sharper delimitation of"Coronophorales", i.e. Nitschkiaceae, leaves it outside. It mimics a diminutiveAcanthonitschkea, but shows a number of alien features, e.g. the paler conicalpapilla of the ascocarps, the structure of the peridium (well shown in Fig. 5d) withrelatively thin cell walls without "Munk pores", the indumentum of hyaline hairsand dark, broad-based spines, the palisade of relatively long asci lining the majorpart of the locule. Our fungus affords a new example that lichenicolous fungi as arule have no really close relatives among fungi inhabiting vascular plants, but it may,20-753873 Svensk Bot. Tidskr. 69 (1975)

322 J. A. Nannfeldtlike the Nitschkiaceae, have Lasiosphaeriaceous ancestors and so it is still justifiedto treat it in this paper.Prof. Santesson has contributed the following descriptions and comments: —"Sometimes very small but characteristic ascocarps of a fungus appear on thediscs of dead, black apothecia of certain species of Sticta. At first, this fungus wastaken for a new species of Acanthonitschkea. The cupulately collapsing ascocarpsarmed with rigid, sharp-pointed spines, the clavate asci tapering to a thin stalk, theabsence of paraphyses, and the small fusiform to ellipsoidal, hyaline spores seemedto indicate a relationship to this otherwise corticolous genus. This suggesion couldnot be upheld. The fungus was found to take a rather isolated position and has to beplaced in a new genus.Nitschkiopsis Nannf. & R. Sant. nov. gen.Perithecia sicca cupulato-collapsa, papilla apicali distincta. Peridium membranaceum,fuscescens. Asci 8-spori, clavati, stipitati, annulo apicali incrassato. Sporae fusiformes velellipsoideae, 1-septatae, hyalinae.—Lichenicola.Typus: N. stictarumNannf. & R. Sant.N. stictarum Nannf. & R. Sant. nov. sp.Perithecia madefacta ± globosa, 0.09-0.12 mm diam., fusco-nigricantes, pilis hyalinis et spinisfuscis ornata. Sporae 6-7x 1.5-2 /xm.—In apotheciis Stictarum.Typus: Kenya, Santesson 22144 f (holotypus: UPS; isotypus: S).The vegetative dark mycelium immersed in the ± dead tissues of the host.Perithecia at first immersed in the necrotic hymenia of the host but later superficial, scatteredor sparsely gregarious, when moist almost spheric with a slightly protruding conical papilla,which is somewhat paler than the rest of the perithecium, when dry collapsed into cup-shapewithout any visible papilla. The upper part of the perithecium furnished with spines and hairs.Spines 1-celled, thick-walled, dark brown to almost black, 12-20 /xm long, ± conical withpointed apex and up to 3-5 /xm broad, basally rather abruptly enlarged to 6-12 /xm. Hairs2-5-celled, thin-walled, hyaline, subcylindrical, 55-85 (xm long and mostly 3.5—1 /xm broad,apically rounded and the apical as well as the basal part slightly enlarged, ca. 5 /¿m and 5-6 /xmrespectively. The dark spines usually 10-25 per perithecium, the hyaline hairs much fewer,usually 3-6 only. Hyaline hairs rarely with dark brown zones and a flattened broad base, thus insome respects transitional to dark spines.Peridium membranaceous, pseudoparenchymous, in thin sections rather pale brown, 10-15/xm thick, the paler papilla ca. 25-30 /xm in diam.Asci 8-sporous, 30-40x5-6 /xm, clavate, rather long-stalked, non-amyloid; apical ring distinct,thickened. No paraphyses visible. Below the papilla a cushion (ca. 15-20 /xm) ofperiphysis-like hyphae.Fig. 5. Nitschkiopsis stictarum (holotypus). (a-c, e) Hairs and spines on the surface of the ascocarps. (a, c)Hyaline hairs with rounded tips and slightly thickened bases, (e) Dark-brown spines with pointed tips andbroad flattened bases. (b) Transitional types with dark parts on the hyaline hairs and broad flattened bases(ca. 700x). — (d,f) Median sections of ascocarps, (d) showing the prominent ostiolarpart (seen also in a),f showing the arrangement of the asci and the periphysis-like hyphae (ca. 500x).Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 323SvenskBot. Tidskr. 69 (1975)

324 J. A. NannfeldtSpores obliquely uniseriate or biseriate, 1-septate, fusiform to ellipsoidal, hyaline, thinwalled,6-7x 1.5-2 p.m.N. stictarum is evidently a parasite killing the attacked parts of the host. Alsoother fungi may be responsible for the necroses and the blackening of the apotheciaand thallus parts as it is not rare to find two or more parasymbionts and parasitessimultaneously attacking a lichen thallus. However, in the Sticta specimens with N.stictarumno other fungus could be identified.Our fungus is very easily overlooked on account of its minute perithecia, whichare as dark as the substrate. When collecting specimens of Sticta lichenologists try toavoid the partly dead specimens in which the fungus grows. However, some samplesof the new fungus were found in lichen herbaria.The hosts of AT. stictarum so far known, viz. Sticta ambavillaria (Ach.) Bory andSt. tomentosa(Sw.) Ach., are two closely related species which seem to have rathersimilar ecological requirements. On the East African mountains the parasite seemsto be common in wet forests and bushlands from 2 500 to 3 500 m s.m.DISTRIBUTION: Most probably widely distributed in tropical regions. At presentknown from East Africa and South America.Africa (all finds on St. ambavillaria): Kenya: Central Prov.: Mt. Kenya, W. slope, NationalPark Road (Naro Moru Track), bamboo zone, alt. 2700-3 100 m, on Arundinaria alpina,23.i. 1970 R. Santesson 22144 f (Type). SW. slope, upper part of bamboo zone, 6.ii. 1922 R. E. &Th. C. E. Fries 4024 b (UPS). E. slope, Urumandi, alt. 3 100 m 1959 M. J. Coe (UPS). — Mt.Aberdare, along Wandera's Track, alt. 3 300 m, on Stoebe kilimandarica, 15.vii. 1971 I. & O.Hedberg 5024 1 (UPS).Tanzania: Arusha Prov.: Mt. Meru, E. slope, road to the crater, alt. 1900 m, on a big tree inthe montane forest, 7.Í.1971 R. Santesson 23073 b (UPS). E. slope, ca. 2 km N. of KitotoCamp, alt. 2450 m, on Stoebe kilimandscharica and Hypericum lanceolatum in dense Ericaceousbushland, 7.i. 1971 R. Santesson 22960, 22969 (UPS). — Kilimanjaro Prov.: Mt. Kilimanjaro,W. slope, E. of Lemosho Glades, Ericaceous zone, alt. 2800-2900 m, on various shrubs,14.i. 1970 R. Santesson 21297 a (UPS).South America (both finds on St. tomentosa): Peru: Dept. Puno: Sachapata, ix.1854 W.Lechler (parasitic on Plantae peruv. 3124; UPS).Brazil: Estado Paraná: Rio Negro 9.x. 1908 P. Dusén (S)."8. The "Nitschkia Group" in EuropeMy studies in the Scandinavian herbaria and in some others have revealed 6 speciesof this group to occur in the Nordic countries and two more in West and SouthEurope, one of which (AT. confertula) should be sought for in south-westernScandinavia. The floristic literature is insufficient to recognize them, and—as shownin Chapt. 2—their nomenclature has been badly confused. Fitzpatrick's wellfoundeddistinction between the two species that had both passed as N. cupularishas astonishingly been neglected by all subsequent European writers.SvenskBot. Tidskr. 69 (1975)

AcanthonitschkeaMiiller and Dennis.Studies in the Coronophorales 4-8 325tristis, which escaped his notion, has been missed also by Arx &N. dissipata Kirschst. is Dermea padi (A. & S. ex Fr.) Fr. (Nannfeldt 1975:62).As Munk (in litt.) has pointed out to me, his Trie ho sphaeria brevispina is in fact amember of this group (cp. above) but his Rostrocoronophora geranii Munk(1953: 96; 1953 b: 87; 1957:282) is a Gnomonia. Gola (1930:192) listed, as present inHb. Saccardo (PAD), a Swedish specimen of an additional species, viz. of theotherwise exclusively North AmericanFracchiaea callistaf =N. callista). On inspectionthe record turned out to be based on a tentative determination by Saccardo of asample sent by Romell (a duplicate of Hb. Romell 16154, S!). Berlese (1900:26)based on the same gathering his Fr. romelliana,which Fitzpatrick (1924: 109) consideredto be a Cryptosphaerellaand which also in my opinion belongs to Coronophoras.lat.Due to the similitude between the different species and the discussions in Chapt. 4of the diagnostic features, no descriptions are given here, only a determinative key.This is followed by detailed accounts of the European distributions so far known ofthe species (for their total distributions see Chapt. 5 and 6). The lists are based on thespecimens revised by me and on reliable literary records. They do not pretend to beexhaustive except for the Nordic countries. Erroneous records have been rectifiedas far as possible.Here, it should be remembered that there exists one not too rare, widely differentPyrenomycete with ascocarps collapsing into cup-shape and thus easily mistaken fora Nitschkia, viz. Sphaeria pomiformis Pers. ex. Fr., later transferred to genera suchas Melanopsamma and Chaeotosphaeria. It is easily known by the ostiolateascocarps (0.25-0.3 mm across), often with some hyaline "setae" (in fact effeteconidiophores) up to 250 pm long, by subcylindrical—clavate asci with a relativelythick and stiff wall, a distinct apical dome and a subapical cushion (both best seenwith Congo Red) and by hyaline, broadly ellipsoidal spores (12-18x5-8 /tm),1-septate with a large drop in each cell.Key to the European species of the "NitschkiaGroup".1. Subiculum with dark stiff bristles, up to 200 p.m long (Acanthonitschkea).Spores 6-9x 1.5-2 pm.Ac. tristis1. No bristles (Nitschkia)2. Asci large (up to 120x25 /im) with >200 spores, arranged in longitudinal rows.Sp. 8-11x1.5-2 pm, suballantoid. Ascocarps 0.3-0.65 mm across, onlyrarely collapsing into cup-shape. N. broomeiana2. Asci much smaller, 8-sporous.3. Ascocarps small (0.2-0.3 mm), caespitose (to gregarious). Sp. 9-13(-16)x2-3(-3.5) fj,m, suballantoid. N. parasitans3. Ascocarps larger (0.3-0.75 mm), gregarious to scattered.4. Part of the peridial cells prolonged into pointed spines (up to 20 p.m long).Sp. 7-14x3^f fim, subcylindrical to narrowly ellipsoidal. N. brevispina20-753873 Svensk Bot. Tidskr. 69 (1975)

326 J. A. Nannfeldt4. External peridial cells rounded or at most mamillate.5. Average spore length 10 /xm.7. Sp. 10-18x2-2.5 /xm, subcylindrical, hyaline. N. cupularis.7. Sp. 12-18x3.5-7 /xm, fusiform, eventually with a very distinct medianseptum and smoky-grey wall. N. collapsa(1) Acanthonitschkea tristis (Pers. ex Fr.) Nannf.Sweden: Skáne: Lund (lignum) E. Fries (Scl. Suec. 386). — Smaland: Kárda, Kállunda,Beech Forest (Fagus periderm) 21.vii.1944 S. Lundell (UPS). — Varmland: Varnum,Niklasdamm (Ulmus inner-bark) vii.1892 H. Hamberg (S). — Uppland: Lohárad, Erken,Bibacken (Sorb. auc. perid., inner-bark and lignum) 19.V.1918 L. Romell 15562 (S, UPS).Solna, Haga (Ulmus inner-bark) 3.Ü.1891 H. Hamberg (S); (Sorb. auc. perid.) 15.vi.1901 L.Romell 16201 (S, UPS). Stockholm between Skuggan and Vartan (Ulmus inner-bark) 5.v. 1888L. Romell 15835 (S, UPS). Uppsala (Bondkyrka) Kvarnbo lund (Corylus av. perid.) 23.V.1926& 2.ÍV.1928 N. Hylander & J.A.N. 4383 (UPS; Weese, Eumyc. sel. exs. 448); Prun. spin.perid.) 24.iv.1928 J.A.N. 4382 (UPS); V&rdsatra Nature Park (Aln. glut, lignum) 9.iv.l928 S.Lundell & J.A.N. 4381 (UPS); (Ulmus inner-bark) 27.iii. 1929 S. Lundell & J.A.N. 2025 (UPS).Arentuna, Storvreta Forest, ca. 4 km SE of the village (Cor. av. perid.) 24.iv.1967 N. Lundqvist5268 (UPS). — Vastmanland: Sala, at the Silver Mine (Cor. av. perid.) 15.viii.1949 R.Morander 1451 (UPS). — Gastrikland: Gavie, Lóvudden (Ulmus fallen decorticated branch)23.iv. 1962 J.A.N. 17118 (UPS).Norway: Akershus: Ó. B as rum at the R. Lysakerelva, above Jar (bark and lignum) 7.VÍ.1962I. Jorstad & J.A.N. 17145b (O, UPS).[Denmark: No material is left relating to the records of N. tristis from Jylland (Larsen1952:13). They may refer to N. grevillii.]Great Britain: Gloucestershire: Saintsbury (Acer) 26.iv.1936 E. W. Mason 1352 & C. G. C.Chesters (IMI 15228, 15230). — Lincolnshire: nr. Brigg (Ac.) 7.iv.l962 G. Morgan-Jones (IMI127 063). — Somerset: Batheaston (perid.) iv.1866 C. E. Broome (Cke, F. brit. 1:269). —Yorkshire: Carperby (Ulmus?) 10.x. 1936 W. G. Bramley (IMI 13490). Howdale (Ac. pseudopl.una cum N. cup. et soc. Eutypa sp.) 3.iii.l957 W. G. Bramley (IMI 68769a); Pickering,Howdale (Ac. pseudopl.) 22.xi.1956 W. G. Bramley (IMI 75384); Pickering, Clunhall Wood(Ilex) 6.i. 1957 W. G. Bramley (IMI 68154c).France: Dép'. Cóte-d'Or: (Juglans r. lignum) F. Fautrey IV: 143 (UPS).Germany: sine loco exactiore (probably Niedersachsen: nr. Gottingen) (Fagus? bark) C. H.Persoon (lectotypus Sph. tristis).Switzerland: sine loco exactiore (Ulmus bark) Morthier 4 (typus Coel. med.). Jura Mts. (Cor.av. perid. and lignum) Morthier (Hb. Barb.-Boiss. 590 p.p.; Fuck., F. rhen. 947 p.p.).Czechoslovakia: Moravia: Hranice (=Mahr.-Weisskirchen) Podhorn (Ulmus fallen decorticatedbranches) 18.vi.1922 F. Petrak (typus Syd. mor.). "Schlesien pr. Gabel" (Fagus perid.and inner-bark) viii.1911 G. v. Niessl. (S).(2) Nitschkia brevispina (Munk) Nannf.Sweden: Uppland:J.A.N. 23901 (UPS).Dalby, "Ormberget" (nr. Grana) (Ulmus) 7.iii.l975 Kerstin Holm &Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 327Denmark: Sjeelland: Hvalsa (Fagus bark, soc. Quaternaria quaternata (vetusta), Nectriasanguinea (vetusta) & Calosphaeria parasitica) 14.ix. 1952 E. Müller, A. Munk & J.A.N,(typus). — Ermelünden (Ulmus) 12.xii.1963 A. Munk (C).Finland: Nyland: Borgâ (Sambucus) ii.1925 W. Nyberg (H).Great Britain: Staffordshire: Oscott College, nr. Birmingham (Ac. ps.pl.) 3.iv.l930 W. B.Groves, E. W. Mason & P. G. M. Rhodes 4481 p.p. (S).(3) Nitschkia broomeiana (Berk.) Nannf.[Denmark: The record from Odense (Salix) E. Rostrup (Lind 1913:195 as Fracch. heterog.)sounds very improbable from a phytogeographical point of view. Munk (in sched. 1945) couldnot find this fungus on the sample (C), nor could I. The very rotten wood is also a most unlikelymatrix. The record should be cancelled.][Belgium: The record from Groenendael (Rosa) by Bommer & Rousseau (1890:244 as Fr.het.) is equally improbable, and the morphological discrepancies noted by them indicate thattheir determination is erroneous.]France: Dép' Vaucluse: Orange (Paliurus acuì.) J. H. Fabre (fide 1878:68).[Three other reputed finds (Dép 1 Haute-Saône: Rigny (Salix) J. Flageolet 37 (PAD); Dép'Rhône: Lyon (Roumeg., F. gall. 5568) and Dép' Seine-Inférieure: Rouen F. Fautrey (UPS) arenot substantiated by the specimens. Fautrey's fungus is N. cupul. s. orig.][Germany: Reported by Kirschstein (1911:289) from Brandenburg (Rathenow) on Robiniapseudacacia. The find was originally published (Kirschstein 1906:51) us Bertiella poly sporanov. gen. et sp. on Quercus. His renaming is definitely erroneous, as seen already from hisillustration. The type specimen (B) shows it to be a unitunicate Pyrenomycete, widely differentfrom the Coronophorales but unknown to me.]Italy: "Lombardia, Veneto, Trentina, Sicilia (Acer camp., Aesculus, Alnus, Buxus, Cercis,Eriobotrya, Lonicera, Rhamnus, Rubus etc.)" fide Traverso (1907:361 as Fr. het.). Severalspecimens seen by me in PAD, S, UPS and W, e.g. Cav., F. longob. 119, and Sacc., Myc. ven.88.Czechoslovakia: Slovakia: Presov (=Epeijes) (Vitis vinif.) vii. 1869 St. Schulzer v. Müggenburg(1870:641-642, Hazslinszky 1873:367; fide Höhnel 1920:130).Yugoslavia: Bosna i Hercegovina: Jajce (=Iaize) (Salix) 1903 Fr.v.Höhnel (fide 1918:139).(4) Nitschkia collapsa (Rom.) Chen.Sweden: Närke: St. Mellösa, Tjuren Isl. (Sorb. auc. inner-bark) 27.vii. 1945 R. Morander 247(UPS). — Uppland: Norrsunda, Rosersberg (S. auc. inner-bark) 7.iv.l890 & v. 1890 J. W.Hamner (S, UPS); (5. auc.? inner-bark and lignum) 7.iv.l890 L. Romell 15563 (S; Romell1892:176). Spânga (S. auc. perid.) 3.vi.l892 H. Hamberg Hb. Romell 15564 (S; Romell1892:177). Stockholm, "ad Holmiam" (S. auc. perid. and inner-bark) 2.vi. 1892 H. Hesselman("sine subie." UPS; "cum subie." S). Uppsala, Slottsbacken (Sorbus sp.? perid. and innerbark)10.v. 1962 N. Lundqvist 3514 (UPS). Uppsala (Bondkyrka) Vârdsâtra (S. auc. perid.,inner-bark and lignum) 12.iv. 1885 L. Romell 15121, 15823 p.p. (S; F. exs. scand. 70 "adUpsaliam", typus; Romell 1889:24); (5. auc. perid.) 31.v. 1885 L. Romell 15823 p.p. (S).Denmark: Sjeelland: 0vered (Fagus bark and "terra nuda") 28.iii.1908 J. Lind (C, S;1913:190 asHerpotr. coll.)[Munk (1953:100; 1957:295) drew evidently his description of "Calyc. coll." from another ofLind's collections, viz. that from Ruderhegn, which is in fact N. grevillii and matches Munk'sdescription. He reported N. collapsa also from Fyn (Skârup) collected by E. Rostrup (asLasiosphaeria spermoides) on Robinia pseudacacia, but I have not seen this collection.]20-753873 Svensk Bot. Tidskr. 69 (1975)

328 J. A. NannfeldtGreat Britain: Evidently not common but widespread, being seen by me (mainly in IMI) fromBedfordshire, Devonshire, Nottinghamshire, Somerset, Yorkshire and the Channel Islands(Guernsey) on Acer, Crataegus, Fraxinus, Prunus spinosa.—As shown by authentical materialfrom Batheaston (W) "Sphaeria tristis var. sporidiis majoribus" (Berkeley & Broome1871:425) is this species.—Fitzpatrick (1923:53) saw in NY a specimen collected by Plowrightin 1873 and labelled Sph. tristis.—The published record of Bertia collapsa from Guernsey (onRibes grossularia and R. rubrum, leg. Marquand) seems to be correct according to the datagiven by Booth (1957:25; cp. Massee 1898:24; Hawley 1923:227).Belgium: Prov. Oost-Vlanderen: Dendermonde (=Termonde) (Ulmus bark) D. Westendorp(Roumeg., F. gall. exs. 3524).France: sine loco exactiore (Berberis, Crataegus, Sorbus) J. E. Chenantais (fide1918:71-73). —Dép' Seine: "prope Parisios" C. H. Persoon ("Sph. tristis", L 910, 269-1008).Montmorency (Crataegus and soil) E. Boudier (typus Trichosph. vag.).(5) Nitschkia confertula (Schw.) Nannf.Great Britain: "On bark &c" (Dennis 1960:212; 1968:334 as Tymp. euomph.).—Evidentlywidespread in England and Wales, seen by me (mainly in IMI) from Cardiganshire, Devonshire,Glamorgan, Gloucestershire, Monmouthshire, Pembrokeshire, Somerset, Surrey andYorkshire (14 localities in all; on Fraxinus associated with Hypoxy Ion rubiginosum) and justreaching Scotland (Wigtownshire: Mull of Galloway, Logon Woods viii,1936C. G. C. Chesters,IMI 13391).Italy: Toscana: "Selva pisana" O. Beccari (typus Coel. becc.).(6) Nitschkia cupularis (Fr. ex Fr.) Karst. s. orig.Sweden: sine loco exactiore E. Fries (Sel. Suec. (ed. 1) 231; typus).Great Britain: Glamorgan: Southerndown (Frax.?, bark) 31.x. 1944 S. J. Hughes 9 (IMI 132617). — Norfolk: King's Lynn (Acer camp.) 13.iii. Ch. E. Plowright (S). Wheatfen Broad(Salix) v. 1946 (IMI 5528). — Yorkshire: Burton, Constable, The Moors (Ulmus) 22.1.1949 J.Webster (IMI 34004; S). Howdale (Frax.) 12.iv.1947 Mr. Rimmington (IMI 14106); (Acerpseudopl.; una cum Ac. trist.) 3.ii.l957 W. G. Bramley 68 (IMI 68769b). Thornton (Crataegus)x. 1945 (IMI 1672).Belgium:Prov. Liège: Malmédy (Frax. ornus, bark) M. A. Libert (Roumeg., F. gall. 1551).France: Dép' Côte-d'Or: Noidan (Carp., lignum) 1890 F. Fautrey (typus Melan. num.) —Dép' Seine-Inférieure: Rouen (Castanea) 9.iii.l890 F. Fautrey 30 (UPS).Germany: Brandenburg: Westhavelland: Gross-Behnitz (Rhamnus cath. perid. and innerbark)6.iv.l898 W. Kirschstein (Typus Herp. rehm.)\ Park Gross-Behnitz (Frax. exc.)12.iv. 1901 W. Kirschstein (B); Klein-Behnitz, Hasellake (Ulmus) x.1905, xii.1909, xi. 1910 &23.xi.1913; (Rosa can.)-, (Coryl. av.) 9.iv.l917; (Prunus spin.) 6.ii.l916 W. Kirschstein (B).Ost-Prignitz: Trignitz i.d. Prignitz (Crat. ox., soc. Diatr. stigma) 25.iii. 1910 O. Jaap 787 (S).Austria: Nieder-Ôsterreich: Wienerwald, Rekawinkel (Quercus) vi.1907 Fr. v. Hôhnel(Rehm, Ascom. 1743).(7) Nitschkia grevillii (Rehm) Nannf.Sweden: Uppland: Uppsala (Bondkyrka) Kvarnbo (Coryl. av.? inner-bark) 19.ix. 1965 N.Lundqvist 4746 (UPS). —Jâmtland: Are, between Are and Sââ (Prunus pad. inner-bark) 1887A. Y. Grevillius (typus).Denmark: Sjœlland: Ruderhegn (Fagus bark) 16.iv. 1909 J. Lind (1913:190 as Herp. coll.; C,Svensk Bot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 329UPS). Rude Skov (Samb. nigr. lignum) 17.ix. 1963 A. Münk (C). Dyrehaven (Fagus? lignum)20.iii. 1965 A. Münk (C).[Larsen's records (1952:13) of N. tristis from Jylland may refer to this species, but nomaterial is left.]Norway: sine loco exactiore M. N. Blytt (O). — Oslo: Ullevold N. Moe (O). —Nordland:Saltdalen (Prunus pad. inner-bark) iii.1824 S. Chr. Sommerfelt (1826:212 as Sph. trist.;Rostrup 1904a: 16 as Coel. trist.; O).Great Britain: Dennis' statement about Calyc. tristis: "On wood normally on that bearingmembers of the Diatrypaceae. Not common" (1960:212; 1968:333) refers p.max.p. to thepresent species. Numerous finds in England. Seen by me (mainly in IMI) from Devon,Durham, Essex, Gloucestershire, Hampshire, Hertfordshire, Norfolk, Oxfordshire, Somerset,Suffolk, Surrey, Wiltshire and Yorkshire; on Acer pseudopl., Aesculus?, Carpinus,Fagus, Ilex, Populus and Ulmus.Belgium: Prov. Liège: Courtrai (Populus inner-bark) D. Westendorp (Herb, crypt. 528).Luxembourg: Kockelscheuer (Frax. bark and lignum) vi. 1901 V. Noppeney (typus Wallr.frax.).France: sine locis exactioribus (Cydonia and Prun. spin.) J. E. Chenantais (fide 1918:64-72as N. trist.). —Dép' Haute-Saône: Rigny (Salix bark) 19.vi. J. Flageolet (typus N. wint.) —Dép' Loire-Inférieur: Nantes "Chêne-Vert" (Populus fast., soc. Eutypa lata) J. E. Chenantais(fide I.e.).Germany: sine loco exactiore (probably Niedersachsen: nr. Göttingen) (Populus trem.?bark) C. H. Persoon (L 910, 270-594) as Sph. cup.; cp. Nannfeldt 1975:53). — Westfalen: Bonn(Aesc. hipp.) Hb. Sehlmeyer (B).Switzerland: Jura Mts. (Corylus av.) Morthier (Fuck., F. rhen 947=Herb. Barb.-Boiss. 590).[Poland: Schroeter's record (1894:313) of N. tristis from Falkenberg (=Niemodiin) belongsprobably here.][Italy: Traverso's records (1907:359) of N. tristis refer probably—at least in part—to N.grevillii, but no specimens have been seen by me.](8) Nitschkia parasitans (Schw.) Nannf.Sweden: Skàne: Lund E. Fries (Sei. Suec. (ed. 2) 231 (UPS; cp. Nannfeldt 1975:54). —Uppland: Dalby, "Ormberget" (nr. Gräna) (Ulmus) 23.ii. 1974 Kerstin & L. Holm 175a (UPS).Soina, Âlkistan (Acer) L. Romell 15668 (S); Ulriksdal (Prun. pad.) 25.ii.1912 & 9.iii.l912 T.Vestergren (S). Stockholm (Sorb, auc.) 1.vi. 1884 L. Romell 15327 (S, UPS); Eriksberg (Tilia)9.iv.l887 L. Romell 15834 (S). Uppsala 1885 L. Romell 15833 (S); (Crataeg.) 5.viii.l884 L.Romell 16053 (S, UPS). —Jämtland: Âre (Ribes rubr.) 2.vii.l930 A. G. Eliasson (S, UPS). —Västerbotten: Umeâ (Prun. pad.) v. 1909 J. Vleugel 113 (1911:332 as N. cup.; C, S).Finland: Nylandia: Borgâ (Caragana, Crat., Sambucus) i.—ii. 1925, v. 1926 & iv.1927 W.Nyberg (H, UPS). Elimä, Mustila (Prun. pad.) 18.vi.l908& 15.vi.l909C. G. Tigerstedt (C,H,UPS). —Regio aboënsis: Âbo (Populus) 9.iv. 1861 P. A. Karsten 1067 (H); (Ulmus) 6.v. 1861 P.A. Karsten 1068 (H; F. fenn. 861). — Tavastia australis: Tammela, Mustiala (Acer, Crat.,Prun. pad. Ribes rubr., Rubus id., Syringa vulg., Tilia) several dates 1866, 1869, 1872 & 1891 P.A. Karsten 1069-1078, 1080-1082 (H, UPS); (Syr. vulg.) 15.v. 1887 Onni Karsten, Hb. Karsten1079(H).Norway: sine locis exactioribus (Ribes rubr., Jug!, ein., Acer plat.) (C, O). — Oslo: M. N.Blytt (0);Bryn (Ribes rubr.) v. 1849 N. Moe (O); Tveita (Aesculus) 1840 M. N. Blytt (O); Töien(Acer plat., Aesc., Caragana arb., Jugl. ein., J. reg., Rhamnus fr.) 1840 & 1849 N. Moe(Rostrup 1904«: 16 as Coel. acervata and C. cup.; C, O). —Oppland: Ringebu (Prun. cer., P.pad., Ribes) S. Chr. Sommerfelt (O).20-753873 Svensk Bot. Tidskr. 69 (1975)

330 J. A. NannfeldtDenmark: Sjmlland: Boserup Skov (Ulmus) 1 .xii. 1964 A. Münk (UPS). —Lolland: Stensgârd("Corylus" =Fagus?) viii.1863 E. Rostrup (Lind 1913:195 as N. cup.; C). —Jylland: Hornslet(Fagus) 26.xii. 1908 0. Winge (C). Ârhus, "Haveselskabets Have" (Acerpseudopl.) 2.11909 0.Winge (C); Risskov (Ac. camp.) 30.xii.1908 0. Winge (C); Marselisborg (Prun. spin.) 9.i.l9090. Winge (C).The Faeroes: Stroma: Sandgaerde (Ribes r.) 2.ix.l938 F. H. Möller (1958:90-91 as N. cup.).[Iceland: Coel. cup. was listed from one locality by Rostrup (1904b: 299) but omitted byLarsen (1932) as he had found only a Mollisia on the relevant specimen (in C). My ownre-examination gave the same result.]Great Britain: Dennis' statement about N. cupularis "On dead branches of Acer, Aesculus,Prunus etc. Not common." (1960:211; 1968:333) refers p.max.p. to the present species.Numerous finds in England. Seen by me (mainly in IMI) from Derbyshire, Essex, Gloucestershire,Hertfordshire, Middlesex, Norfolk, Northamptonshire, Staffordshire, Surrey and Yorkshire,on Acer pseudopl., Aesculus, Fagus, Malus, Tilia, and Ulmus.Belgium: Prov. Liège: Malmédy (Acer ps.pl.) M. A. Libert (Roumeg., F. gall. 1488; Thum.,Myc. univ. 1947).France: sine loco exactiore (Aesc.) L. Dufour 611 (Hb. Fries UPS). — Probably Dép'Bas-Rhin: nr. Strasbourg and/or Dép* Vosges: nr. Bruyères-en-Vosges (Alnus, Carp., Jugl. &Robinia pseudac.) (Moug. & Nestl., St. crypt, vog.-rhen. 771 & 1455). — Dép' s Seine &Seine-et-Oise: common at Paris and Versailles (Acerps.pl., Aesc. h„ Juglans r.,Populus nigra,Ribes rubr., Sorbus hybr., Ulmus etc.) Tulasne (1865:82; 1931:77; as Sph. cup.). — Dép'Seine-Inférieure: Rouen (Tilia) A. Malbranche 512 (typus Coel. anc.). —Dép' Vienne: Saint-Romain-sur-Vienne (Rob. ps.) S. DeLacroix (Desm., Pl. crypt. 11:780).Germany: sine loco exactiore G. Kunze (Hb. Fries UPS; "Sph. cupularis" det. Kze). —"Germania" (probably Nedersachsen: nr. Göttingen) (Crat.) C. H. Persoon (L 910, 270-591 asSph. cup. cp. Nannfeldt 1975:53. — Westfalen: Bonn (Aesc. h.) Hb. Sehlmeyer (B). —Hessen: Ostrich (Acer, Prun. mahaleb, Carp.) L. Fuckel (F. rhen. 968; Hb. Barb.-Boiss. 591).— Bayern: Oberbayern: Grosshesselohe (Acer ps.pl.) iv.1892 J. N. Schnabl (S).Brandenburg: Berlin Botanic Garden (Ulmus carp. f. suberosa) iv.-vi. 1888 P. Sydow (Myc.march. 1915; Rehm, Ascom. 983). Stegliz (Tilia) xi.1894 P. Sydow (Myc. march. 2340). CharlottenburgSchlossgarten (Aesc. h.)viii. 1894 P. Sydow (Myc. march. 4131). —Sachsen: Leipzig(Rhamnus) vi. 1874 G. Winter (S).Switzerland: Ct. Bern: Bern (Aesc. h.) autumn G. Otth (typus Nectriae hipp.). — Ct. Ticino(fide Traverso 1907:359 as N. cup.).Austria: Niederösterreich: Pfaffenberg nr. Deutsch-Altenburg (Crat. ox.) xi.1940 F. Petrak(Myc. gen. 177). Sonntagberg (Aesc. h.) x.1914 P. Strasser (1915:95; W). — Oberösterreich:Vöcklabruck (Pr. spin.) vii. 1948 and Wels (Ulmus) vii. 1968 K. W. Kuchar 1974:176 as N. cup).Poland: Warszawa, Mokotöw (Ulmus) 17.x. 1962 J. Kochman (Myc. pol. 326). — Schroeter'srecord (1894:313 as N. cupularis) from Wroclaw (=Breslau) belongs certainly here.Czechoslovakia: Moravia: Hranice (=Mähr.-Weisskirchen) Podhorn (Ulmus) 9.x. 1922 F.Petrak (Fl. boh.-mor. 11:1:1634). Mähr.-Schönberg (Aesc. h.) l.iv. 1878 J. Paul (typus N.morav.). Sternberg (Ulmus?) iii. 1926 J. Piskor (fide Petrak 1927:362 asN. cup.).Italy: Traverso's paragraph about N. cupularis: "Piemonte, Veneto, Canton Ticino,Trentino, Lazio, Napoletano, Sicilia (Acer, Carpinus, Corylus, Fraxinus, Gleditschia,Mespilus, Prunus, Syringa, Tilia, Ulmus etc.)" (1907:359) refers—at least p.max.p.—to thepresent species. Several specimens seen by me, e.g. Sacc., Myc. ven. 1449.Soviet Union (European part): "Prov. Nowgorod: Bologue" (Ulmus mont.) M-vii. 1897 W.26Tranzschel (S).SvenskBot. Tidskr. 69 (1975)

AcknowledgementsStudies in the Coronophorales 4-8 331The present studies.could not have been performed without the kind and generous support ofnumerous institutes and individuals, too many to be fully specified here. My thanks are thusdue to the directors and staffs of the following herbaria from which relevant material has beenreceived on loan, viz. B, C, COI, FH, H, IMI, L, LD, LPS, P, PAD, PC, S, and W. I am furthergreatly indebted to Drs L. Holm and N. Lundqvist (Uppsala), Prof. A. Munk (Copenhagen)and Prof. R. Santesson (Stockholm) for valuable information, good advice and stimulatingdiscussions, and to Prof. Santesson also for contributing the description of his new lichenicolousspecies.Referencesadditional to those listed in the first instalment (Nannfeldt 1975)Ainsworth, G. C. & Bisby, G. R. 1943. A Dictionary of the Fungi. London.— 1971. Id. Sixth ed. London.Berkeley, M. J. 1875. Notices of North American Fungi (contd.). Grevillea 4 (=n. 30): 45-52("1876").— 18766. Id. Ibid. 4 (=n. 32): 141-162.Berkeley, M. J. & Broome, C. E. 1871. Notices of British Fungi. Ann. Mag. Nat. Hist.4(7): 425-436.Berlese, A. N. 1888. Fungi moricolae 5. Padova. (n.v.)— 1900. Icones fungorum 3. Patavii.Berlese, A. N. & Peglion, V. 1892. Micromiceti toscani. . . N. Giorn. Bot. Ital. 24(3): 97-172.Bommer, E. & Rousseau, M. 1890. Contributions à la flore mycologique de Belgique. Bull.Soc. Roy. Bot. Belgique 29(1): 206-302.Booth, C. 1957. Studies of Pyrenomycetes 1 ... 2. Melanopsamma pomiformis and itsStachybotrys conidia. Mycol. Pap. (C.M.I.) 68.Boudier, E. 1910 a. Icones mycologicae .. .3. Plances 422 à 600. Paris. (" 1905-1910")— 1910 b. Id. 4. Texte descriptif. Paris. ("1905-1910")Bresadola, G. 1914. In: De Wildeman, E., Additions à la flore du Congo. Bull. Jard. Bot.Bruxelles 4(1): 3-241.Cain, R. F. 1972. Evolution of the Fungi. Mycologia 64(1): 1-14.Cash, Edith K. & Davidson, R. W. 1940. Some new species of Ascomycetes on coniferoushosts. Ibid. 32(6): 728-735.Chadefaud, M. 1960. Les végéteaux non vasculaires (Cryptogamie). In: Chadefaud, M. &Emberger, L., Traité de Botanique systématique 2. Paris.Chardon, C. E. et al. 1940. Ascomycetes from the State of Minas Geraes (Brazil). Mycologia32(2): 172-204.Chenantais, J. E. 1918. Études sur les Pyrénomycètes. Bull. Soc. Mycol. France34 (1-2): 47-73.Chesters, C. G. C. 1939. The Chipping Campden Foray, 27-31 May 1938. Trans. Brit. Mycol.Soc. 23(3): 235-238.Cooke, M. C. 1878. Ravenel's American Fungi (contd.). Grevillea 7 ( =n. 42): 43-54.— 1887. Synopsis Pyrenomycetum (contd.). Ibid. 15(=n. 75):80-86.— 1892. Additional Fungi Descriptions. Ibid. 20 (=n. 96): 106-107.Cooke, M. C. & Ellis, J. B. 1877. New Jersey Fungi (contd.) Ibid. 5(=n. 35): 89-95.20-753873 Svensk Bot. Tidskr. 69 (1975)

332 J. A. NannfeldtDennis, R. W. G. 1960. British Cup Fungi and their allies . . . Dorking (Ray Society).— 1968.British Ascomycetes. Stuttgart.Doidge, Ethel M. 1950. The South African Fungi and Lichens to the end of 1945. Bothalia5:1-1094.Donk, M. A. 1974. Check List of European Polypores. Verh. K. Nederl. Akad.Afd. Natuurk. 2(62).Wetensch.,Ellis, J. B. & Everhart, B. M. 1890. New North American Fungi. Proc. Akad. Nat. Sei.Philadelphia 1890:219-249.— 1892. The North American Pyrenomycetes . .. Vineland, N. J.Fabre, H. 1878. Essai sur les Sphériacées du département de Vaucluse. Ann. Sei. Nat. Bot.6(9): 66-118.— 1883. Essai sur les Sphériacées du dép. Vaucluse. Ibid. 6(15): 31—69.Fairman, Ch. E. 1921. The fungi of our common nuts and pits. Proc. Rochester Acad. Sei.6(3): 73-115.Feltgen, J. 1905. Vorstudien zu einer Pilz-Flora des Grossherzogthums Luxemburg. 1.Ascomycetes. Nachträge 3. Ree. Mém. Trav. Soc. Bot. Luxembourg 16: 1-328. ("1902-1903")Fries, E. 1830. Ecloge fungorum . . . Linnaea 5:497-553.Fuckel, L. 1871. Symbolae mycologicae . . . Erster Nachtrag. Jb. Nassau. Ver. Naturkunde25-26:287-347.Gola, G. 1930. L'Erbario micologico di P. A. Saccardo. Catalogo: Atti Acc. Sei. Ven. Trent.Istr. 21, Suppl. 1.Griffith, H. Br. 1973. Fine structure of seven unitunicate pyrenomycete asci. Trans. Brit.Mycol. Soc. 60 (2): 261-21 \.Hansford, C. G. 1956. Australian Fungi 3. New species and revisions (contd.). Proc. Linn. Soc.New South Wales 81 (I): 23-51. ("1957")Hawley, H. C. 1923. Notes on some British Pyrenomycetes. Trans. Brit. Mycol. Soc.8(4): 226-230.Hazslinszky, Fr. 1873. Einige neue oder wenig bekannte Arten der Pilzflora des südöstlichenUngarns. Verh. zool.-bot. Ges. Wien 23:361-368.Hennings, P. 1895. Fungi. In: Engler, A., Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete.In: Deutsch Ost-Afrika 5 C: 30-61.— 1897. Beiträge zur Pilzflora Südamerikas 2. Hedwigia 36(3^f): 190-246.— 1898. Über verschiedene neue und interessante märkische Pilzarten, besonders aus derUmgebung von Rathenow. Verh. Bot. Ver. Prov. Brandenburg 40: xxv-xxx.— 1900. Fungi 2. In: Warburg, O., Monsunia . . . 1: 137-174.— 1908. Fungi S. Paulenses 4 a cl. Puttemans collecti. Hedwigia 48(1): 1-20. ("1909")Höhnel, F. v. 19066. Fragmente zur Mykologie (2. Mitteilung, Nr. 64 bis 91). Sitz. ber. Akad.Wiss. Wien, Math.-nat. Kl. 1(115): 649-695.— 19096. Id. (7. Mitteilung, Nr 289 bis 353). Ibid. 1(118):813-904.Hopkins, J. C. F. 1938. A preliminary list of Rhodesian Fungi. Trans. Rhodesian Sei. Ass.35(2): 97-127.— 1939. A descriptive list of plant diseases in Southern Rhodesia (and their control). Mem.Dept. Agric. Southern Rhodesia 2.Hughes, S. J. 1953. Fungi from the Gold Coast 2. Mycol. Pap. (C.M.I.) 50.Karsten, P. A. 1879. Quaedam ad mycologiam addenda. Medd. Soc. F. Fl. Fenn. 5:38^12.Kickx fils, J. 1835. Flore cryptogamique des environs de Louvain. (n.v.)Kirschstein, W. 1906. Neue märkische Ascomyceten. Abh. Bot. Ver. Prov. Brandenburg48:39-61.SvenskBot. Tidskr. 69 (1975)

Studies in the Coronophorales 4-8 333— 1939. Über neue, seltene und kritische Ascomyceten und Fungi imperfecti. Ann. Mycol.37(1-2): 88-140.Kuchar, K. W. 1974. Beitrag zur Kenntnis der oberösterreichischen Pilzflora. Sydowia26: 171-183.("1972")Kuntze, O. 1898. Revisio generum plantarum . . . 3(2). Leipzig.Larsen, P. 1932. Fungi of Iceland. The Botany of Iceland 2(3): 449-607. ("1931")— 1952. Studies in Danish Pyrenomycetes. Dansk Bot. Ark. 14(7).Lind, J. 1913. Danish Fungi. . . Copenhagen.Lindau, G. 1897. Pyrenomycetineae. In: Engler & Prantl, Die natürlichen Pflanzenfamilien1(1): 321—491.Martens, P. 1937. Les Ascomycètes à asques polyspores. Bull. Soc. Mycol. France52 (4): 379-407.Mason, E. W. 1940. Presidential address on specimens, species and names. Trans. Brit. Mycol.Soc. 24(2): 115-125.Massalongo, C. 1889. Contribuzione alla Mycologia veronense. Mem. Accad. Agric. ArtiComm. Verona 65(2): 131-283.— 1902. Novitates florae mycologicae veronensis. Ibid. 78:81-163.[Massee, G.] 1898. British Mycology. Trans. Brit. Mycol. Soc. 1 (1):20-24.Miller, J. H. 1942. South African Xylariaceae. Bothalia 4(2): 251-272.Möller, F. H. 1958. Fungi of the Faeröes 2. Copenhagen.Montagne, C. 1840. Seconde centurie de Plantes cellulaires exotiques nouvelles. Ann. Sci.Nat., Bot. 2 (14): 321-350.— 1856. Sylloge generum specierumque Cryptogamarum . . . Parisiis.Müller, E. & Booth, C. 1972. Generic position of Sphaeria phaeostroma. Trans. Brit. Mycol.Soc. 58(1): 73-77.Mundkur, B. B. & Ahmad, S. 1946. Revisions of and additions to Indian Fungi 2. Mycol. Pap.(C.M.I.) 18.Münk, A. 19536. The System of the Pyrenomycetes . . . Additions and corrections. Bot.Tidsskr. 50(1): 86-90.— 1962. An approach to an analysis of taxonomic method with main reference to higher fungi.Taxon 11(6): 185-190.Nannfeldt, J. A. 1972. Camarops Karst. (Sphaeriales-Boliniaceae) with special regard to itsEuropean species. Svensk Bot. Tidskr. 66 (4): 335-376.— 1975. Stray studies in the Coronophorales (Pyrenomycetes) 1-3. Ibid. 69(l):49-66.Otth, G. 1869. Sechster Nachtrag zu dem in Nr. 15-23 der Mittheilungen enthaltenen Verzeichnisseschweizerischer Pilze. Mitth. naturf. Ges. Bern 1868:37-70.Ou, S. H. 1936. Supplement notes on Ascomycetes from China 2. Sinensia 7(2): 194-211.Parguey-Leduc, Agnès. 1966. Recherches sur l'ontogénie et l'anatomie comparée desascocarpes des Pyrénomycètes ascoloculaires 2. Les ascocarpes des Pyrénomycètesascoloculaires unituniqués. Ann. Sci. Nat. Bot. 12 (8): 1-110. ("1967") Also diss., Paris.— 1968. Recherches préliminaires sur l'ontogenie et l'anatomie comparée des ascocarpes desPyrénomycètes ascohyméniaux 3. Les asques des Sordariales et leur ascothécies, du type"Diaporthe". Rev. Mycol. 32 (5): 369^107. ("1967")Patouillard, N. 1889. Fragments mycologiques. Champignons extra-européens. Journ. de Bot.3(10): 165-168.— 1921. Quelques champignons du Tonkin (suite). Bull. Soc. Mycol. France 40 (1): 28-37.Patouillard, N. & Gaillard, A. 1889. Champignons du Vénézuéla et principalement de la régiondu Haut-Orénoque, récoltés en 1887 par M. A. Gaillard (suite). Ibid. 4(3): 92-129.Paul, J. 1909. Beitrag zur Pilzflora von Mähren. Verh. naturf. Ver. Brünn 47:119-148. (" 1908")20-753873 Svensk Bot. Tidskr. 69 (1975)

334 J. A. NannfeldtPenzig, O. & Saccardo, P. A. 1897. Diagnoses fungorum novorum in insula Java collectorurn.Series prima. Malpighia 11 (4): 387-409.— 1904. Icones fungorum javanicorum. Leiden.Petch, T. 19176. Additions to Ceylon Fungi. Ann. R. Bot. Gard. Peradeniya 6(3): 195-256.Petrak, F. 1923. Mykologische Notizen 6. Ann. Mycol. 21 (3^4): 182-335.— 1927. Beiträge zur Pilzflora von Sternberg in Mähren 2. Ibid. 25 (3-4): 344-388.— 1950. Über die Gattung Gaillardiella Pat. Sydowia 4:158-162.— 1952 a. Die Gattung Schizocapnodium. Ibid. 6(1-4): 265-267.— 19526. Über die Gattungen Scortechinia Sacc. und Teratonema Syd. Ibid. 6 (5-6): 396-398.— 1952c. Beiträge zur Pilzflora von Florida. Ibid. 6 (5-6): 399-406.— 1952d. Biciliospora n. gen., eine neue Gattung der Sphaeriales. Ibid. 6(5-6): 429-432.— 1953. Ein Beitrag zur Pilzflora Floridas. Ibid. 7(1-4): 103-116.— 1962a. Ergebnisse einer Revision der Grundtypen verschiedener Gattungen der Askomyzetenund Fungi imperfecti (115-125). Ibid. 16:353-361.— 19626. Über die Gattung Cryptosphaerella Sacc. Ibid. 16:362-367.Plowright, Ch. B. 1876. Californian Fungi. Collected by H. W. Harkness, M.D. Grevillea5(=n.34):14.Rehm, H. 1903. Ascomyceten-Studien 1 .Hedwigia 42, Beibl. 4: (172)-(176).— 1907. Ascomycetes exs. Fase. 40. Ann. Mycol. 5(6):465-473.Romell, L. 1889. Fungi aliquot novi, in Suecia media et meridionali lecti. Bot. Not.1889(1): 23-26.— 1892. Nägra ord om Sphaeria astroidea ... och Bertia collapsa. Ibid. 1892 (4): 170-178.Rostrup, E. 1904a. Norske Ascomyceter. Vidensk.-Selsk. (Christiania) Skr. 1. Math.-nat. Kl.1904 (4).— 19046. Islands Svampe. Boi. Tidsskr. 25(3):281-335.Roumeguère, C. & Saccardo, P. A. 1881. Reliquiae mycologicae Libertianae. Series altera.Rev. Mycol. 3 e année

Studies in the Coronophorales 4-8 335Traverso, J. B. 1907. Pyrenomycetae. Fase. 2. In: Flora Italica Cryptogama 1: Fungi. RoccaS. Cassiano.Vleugel, J. 1911. Zweiter Beitrag zur Kenntnis der Pilzflora in der Umgebung von Umeâ.Svensk Bot. Tidskr. 5(3): 325-350.Wallroth, F. G. 1833. Flora cryptogamica Germaniae 2. Norimbergae.Wehmeyer, L. E. 1926. A biologie and phylogenetic study of the stromatic Sphaeriales. Amer.J. Bot. 13 (12): 575-645.Werdermann, E. 1923. Über die Gattung Teratonema Syd. Ann. Mycol. 21 (3-4): 336-339.Winter, G. 1883. Fungi nonnulli novi. Hedwigia 22(1): 1-3.Zeller, S. M. 1927. Contributions to our knowledge of Oregon fungi 2. Mycological notes for1925. Mycologia 19(3): 130-143.Additional NoteSivanesan (Trans. Brit. Mycol. Soc. 64(3): 441-445, 1975) has recently transferred Lasiosphaerianoonae-daniae (discussed in Chapt. 4) to his new, two-species genus Lasiosphaeriella,which he placed in Lasiosphaeriaceae sensu Lundq. and characterized by ostiolate ascocarps,asci without any apical apparatus, and spores surrounded by a mucilaginous sheath andgerminating by phialides. The type species is L. dennisii Sivan. from Uganda. Both specieshave tuberculate ascocarps and "Munk pores". In both the cells of the basal part of theperidium tend to lie in radiating rows.20-753873 Svensk Bot. Tidskr. 69 (1975

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