Allophylaria subliciformis (Pezizellaceae, ), a rare species rediscovered in Europe and North America

Rubén MARTÍNEZ–GIL Abstract: the macro- and microscopic characters of the very rarely recorded Allophylaria subliciformis, type Hans-Otto BARAL species of Allophylaria, are described and illustrated from fresh collections made in spain and Germany. a sample from Canada is also reported. a history of the genus Allophylaria is given. new data, ecological and chorological information and some comments on the confusion with Crocicreas and Cyathicula are added. rDna sequences of ssU, its, and lsU were gained which support placement of the genus in the Pezizella- Ascomycete.org, 10 (2) : 67–75 ceae. Mise en ligne le 22/04/2018 Keywords: , chorology, fungi, hemiamyloid, . 10.25664/ART-0230

Introduction cicreas subhyalinum (rehm) s.e. Carp., a species which was found by baral (in baral & KrieGlsteiner, 1985) to show the typical characters of Allophylaria: a hemiamyloid reaction of the “Calycina-”(Laetinae- Karsten (1869: 103) proposed a section Allophylaria within the via-) like ascus apical ring, the absence of crystals, and cylindrical genus Peziza Dill. ex Fr., in which he placed on pp. 147–148 two vacuolar bodies in paraphyses and terminal cells of excipulum. species, Peziza eucrita P. Karst. and the newly described P.‘sublicoides’ arenDholz (1989: 288) reviewed the nomenclature and history of P. Karst. (renamed to A. ‘sublicaeformis’ in the index on p. 205). the Allophylaria in detail. also he accepted nannfeldt’s view by recog- very same sectional diagnosis was reproduced in a “repertorium” by nizing Allophylaria as an independent genus, but he did not present rabenhorst (1869), who named Karsten’s 25 sections “Untergattun- a concept of included species. gen” (subgenera). a year later, Karsten (1870: 243) raised this section baral (in baral & KrieGlsteiner, 1985: 94) included A. subhyalina to generic rank and included four species: Allophylaria eucrita (rehm) baral in Allophylaria and defined the genus by an ectal ex- (P.Karst.) P. Karst., A.‘sublicaeformis’ (P. Karst.) P. Karst., A. clavuliformis cipulum of textura oblita that may get thin-walled with age and does (P. Karst.) P. Karst. and A. byssacea (P. Karst.) P. Karst. only one more not form triangular scale-like structures at the margin, by the mostly year later, Karsten (1871: 164) reduced Allophylaria to synonymy with red- but never blue-reacting apical ring (in lugol, erroneously as Pezicula tul. & C. tul. Karsten (1885: 130) reinstalled the genus Allo- phylaria with the new synonym Helotidium sacc. (type H. nanum “Melzer’s reagent” and “dextrinoid”), and cylindrical, refractive vac- sacc.), by including A. ‘sublicaeformis’, A. clavuliformis, A. byssacea, uolar bodies (as “lipid bodies”). he suggested a relationship of Ca- and A. phyllophila (Desm.) P. Karst., but he excluded A. eucrita which lycina (= Pezizella) with the nannf. because of the he retained in Pezicula. the latter species is considered a typical Pez- similar type of apical ring that occurs in various genera of that family, icula up to now, whereas A. phyllophila was transferred to Calycina including members today recognized in Lachnaceae raitv., but re- tained Allophylaria in the (as ). nees ex Gray in baral & KrieGlsteiner (1985). baral (1987a) presented an overview on the apical ring types in Allophylaria was synonymized with Hyaloscypha boud. by boUDier (1907), a view which was subsequently adopted by Grelet (1951), the Helotiales and classified Allophylaria (figs 21–22) as Laetinaevia although the two genera differ by glabrous vs. hairy apothecia. type, which was also observed in Ascocoryne, Hysteropezizella, and nannFelDt (1932) slightly altered Karsten’s concept and enlarged Heterosphaeria (figs 18–20), in contrast to the Hymenoscyphus type the genus by including various further species with more or less in Cyathicula (fig. 12). hairless apothecia, for instance, Allophylaria macrospora triebel & baral (1996) confirmed the Hymenoscyphus type for (W. Kirschst.) nannf., and at least one typical member of Calycina, three species of Cyathicula (figs 8–12) but found a Conchatium type C. vulgaris (Fr.) baral (= Pezizella sordida (Fuckel) Fuckel), the type (Calycina-like) in four other species of that genus (figs 4–7), which species of Pezizella Fuckel, as well as a member of Psilachnum höhn., appeared to form a transition to the Calycina type in Crocicreas Fr. Psilachnum chrysostigma (Fr.) raitv. nannFelDt (op. cit.) proposed Pez- s.str. For this reason, they hesitated to distinguish between Croci- iza‘sublicoides’ as lectotype of Allophylaria and placed the genus in creas and Cyathicula. they recommend that reliable taxonomic the family Helotiaceae rehm. although he considered Allophylaria study in the Helotiales should be performed using fresh material, and Pezizella as synonyms, he adopted Allophylaria because Pezizella since some characters, such as the presence and shape of the vac- was interpreted in different ways. both genera were firstly recog- uolar bodies, are only visible in living cells of the paraphyses and ex- nized at the generic level in 1870, although nannfeldt asserted that cipular hyphae. these structures are taxonomically relevant, for Allophylaria was a year younger than Pezizella. example, to distinguish Allophylaria from both Cyathicula De not. Dennis (1956) accepted Allophylaria as a small group of species and Crocicreas Fr.: Allophylaria always has large and elongated vac- distinct from Pezizella, with a “phialeoid or pezizelloid excipulum, uolar bodies that occupy almost all the space of the terminal cell, non-lanceolate paraphyses, and relatively large asci and spores”. he whereas those of Cyathicula are small, globose, and in high number redescribed the type of A. ‘sublicoides’ but mentioned also a record (multiguttulate) (baral & KrieGlsteiner, 1985; baral, 1992). Crocicreas on Chamaenerium angustifolium made by nannfeldt in sweden. in its narrow circumscription lacks Vbs at all (n.V. Filippova, pers. KorF (1971) recombined A.‘sublicoides’ in the genus Cyathicula De comm.; see baral, s.d./b). triebel & baral (op. cit.) also commented not., probably because of the phialeoid ectal excipulum in both on other important characters of the genus Allophylaria, such as the genera. a decade later, CarPenter (1981) synonymized Cyathicula consistent absence of crystals on the apothecial outside, the fre- with Crocicreas, but retained Allophylaria as a distinct genus, due to quently substipitate ascocarps, and the apical ring reacting red- differences in excipular structure (unbranched and very parallel hy- brown in the presence of iKi and having an obconical shape of the phae in Allophylaria). however, he proposed the combination Cro- Laetinaevia type (similar to Calycina).

67 hUhtinen (1990: 50) considered boudier’s synonymization of Allo- curvulae, longit. 15–26 mmm., crassit. 6–7 mmm., guttulis 2 magnis phylaria with Hyaloscypha as a mystery, but baral (2015) questioned praeditae; thecae clavatae, longit. 160–170 mmm., crassit. 11–13 the consistent presence of hairs in Hyaloscypha by including also a mmm.; paraphyses graciles, crassit. circiter 1,5 mmm., apicem versus hairless member in that genus, H. minuta (spooner & Dennis) baral. leniter incrassatulae. baral & räMä (2015) and baral (in JaKlitsCh et al., 2016) resurrected a Ad caules Artemisiae vulgaris dejectos exsiccatos exeunte mense Oc- separate family Pezizellaceae Velen. to accomodate genera such as tobri in Mustiala lecta. Calycina and Allophylaria and many others, which all have this type of apical ring in combination with refractive vacuolar bodies (Vbs) Holotype (fide hUhtinen, 1990: 242): [Finland, tavastia australis,] in the paraphyses and cortical cells of the ectal excipulum, the latter Mustiala, ad Art.[emisia], 31.X.1867, P.a. Karsten (h, herb. Karsten often showing gelatinized cell walls. 884, not examined). the present molecular data of the type species of Allophylaria support placement in the family Pezizellaceae near Calycina, partic- Macroscopic description ularly near C. citrina (hedw.) Gray. Ascocarps (0.15–)0.25–0.5 mm in diam., gregarious in large num- bers, whitish to pale yellow, goblet–shaped, and with a small cylin- Materials and methods drical stipe of 0.1-0.2 mm height and 0.1-0.15(–0.17) mm width, pale yellow- to grey-brown in the upper part and darker brown below, superficial. the collections presented here have been photographed macro- and microscopically. For this purpose, a sony α68 Dslr camera and Microscopic description a Minolta 100 macro lens have been used on a tripod under natural Asci *(140–)150–190(–240) × (15–)15.5–17.5(–19) μm {5}, †135– light (r.M), or a nikon Coolpix 4500 on a tripod in combination with 170 × (10–)11–14(–15) µm {3}, cylindric-clavate, narrowed towards a macro lens (russian horizon lens 10×) at maximum zoom (32 mm), the base, apex with strongly hemiamyloid apical ring of Laetinaevia illuminated by 4 miniature leDs (h.b.). type {6}, blue at low and red at high iKi concentration (rb), or some- For the microscopic observation and its corresponding descrip- times almost directly red (rr), negative in Mlz but deep blue in iKi tion, a Motic DM–ba 200 optical microscope was used, with a Mot- or Mlz when Koh-pretreated, with croziers {4}, 8-spored, spores icam 2000 micro-camera connected to a computer and the official obliquely biseriate, pars sporifera *58–65 µm. Ascospores *(19.5–) “Motic images Plus 2.0” program (r.M.), or a Coolpix 4500 adapted 21–26(–30) × (6–)6.5–7(–7.5) μm {6} (Q = 3.1–4.1, n = 68, r.M.), †(17–) free-hand to a Kpl 10×/20 zeiss wide field ocular and 100× zeiss 19–23(–24) × 5–6.3 µm {2}, cylindric-ellipsoid, ends obtuse to almost achromatic objective (h.b.). subsequently, the photos were treated subacute, homopolar, inequilateral to mostly slightly (to medium) and arranged with computer programs (adobe Photoshop, aCD- curved (allantoid), usually non-septate within living asci but some- see). times also 1-septate {1}; overmature with 1–3 septa, scarcely con- both macro- and microscopic data were gained from the fresh stricted at the septa; surface pale lilac in Crb due to a thin gel layer; specimens, but some details were examined from the conserved lipid bodies (2–)3–4(–4.5) µm diam., 1(–3) in each half, surrounded herbarium material. For the measurement of spores, only those that by many small ones (lipid content 4.5–5); in septate spores with one were recently discharged from the asci were measured. lb of 4–5.5 µm diam. in each cell; freshly ejected spores covered the mountants and reagents used for the microscopic observa- with a delicate sheath that soon detaches and is then difficult to see; tion were h o (tap water), lugol’s solution (iKi, with a concentration 2 sometimes forming chalara-like phialides on the spores. Paraphy- of ~1% i , added to the water mount), Cresyl blue (Crb, aqueous), 2 ses filiform, slightly thickened at the cylindric-clavate apex to and Cr (Congo red, aqueous). iKi was particularly used for obser- *(2–)3–5(–5.7) μm, often somewhat curved, rarely antler-like, sep- vating the hemiamyloid reaction of the ascus apical ring, and Crb tate, terminal cell *(28–)32–41(–44) µm in length, containing one or for staining gel on the spore wall. a few large, hyaline to pale yellowish vacuolar bodies that occupy Voucher specimens were deposited in the private herbaria of almost all its space, reaching 10–20 µm from top, Vbs disintegrating rubén Martínez (r.M.) (at the headquarters of the Verpa Cultural into small, orange-yellow to red-brown drops in the presence of iKi. Group Mycological of logroño), adrian Carter (a.C.), and h.o. baral Medullary excipulum hyaline, formed of long cylindrical cells 2– (h.b.). rDna sequences of Allophylaria subliciformis were deposited 4μm diam. Ectal excipulum in receptacle hyaline, of textura por- in Genbank. “s.d.” means undated. recta to porrecta-oblita, composed of cylindrical cells in parallel and unbranched, partly undulating disposition, cells *3–8 μm wide, Taxonomy moderately thick-walled (strongly so in dead state), more thin- walled with age, agglutinated by a gelatinous outer wall layer that Allophylaria subliciformis (P. Karst.) P. Karst. [as ‘sublicaeformis’], swells in dead state; margin formed by chains of cylindrical cells, ter- Not. Sällsk. Fauna Fl. Fenn. Förh., 11: 243 (1870). minal cells cylindrical to claviform, *12–24 × 4–8 μm, containing Vbs similar to the paraphyses; stipe brown (in Koh olive-brown), some- ≡ Peziza subliciformis P. Karst. [as ‘sublicoides’], Not. Sällsk. Fauna Fl. Fenn. Förh., 10: 148 (1869) – in index changed to ‘sublicaeformis’. times with yellowish exudate. ≡ Pezicula subliciformis (P. Karst.) P. Karst., Bidr. Känn. Finl. Nat. Folk (Mycol. Fenn.), 19: 164 (1871). Ecology: ruderal flora of gardens and recultivated pits, exposed ≡ Hyaloscypha subliciformis (P. Karst.) boud., Hist. class. Discom. Eur.: to the sun, on previous year’s fallen stems or sometimes husks lying 127 (1907). more or less moist among low grasses, of Oenothera biennis {1}, ≡ Cyathicula subliciformis (P. Karst.) Korf [as ‘sublicoides’], Phytolo- Melilotus sp. {1}, S. canadensis {1}, indet. herbaceous plant {2}. assoc.: gia, 21(4): 203 (1971). Calycina herbarum {1}, Exarmidium inclusum {1}, Hohenbuehelia sp. {1}, Orbilia vitalbae {1}, Unguiculella eurotioides {1}. Phenology: Xi–i. Taxonomic position: Pezizellaceae, Helotiales, Leotiomycetidae, , , ascomycota, Fungi. Specimens included : SPAIN. Community of La Rioja, 14 km sW of logroño, 0.2 km se of hornos de Moncalvillo, 42° 23’ 22’’ n – 2° Original description (Karsten 1869) 35’ 1’’ W, 683 m alt., on unidentified herbaceous dicot stem, Apothecia gregaria, exigua, crassa, altit. circiter 0,3 mm., cupula 3.Xii.2016. leg. l. ballester, herb. r.M. 2374, Genbank Mh221035 – lutescente, latit. circiter 0,1 mm., cum stipite brevissimo crasso pallido GERMANY. Mecklenburg–Vorpommern, 20 km WnW of schwerin, vel lutescenti–pallido obconica; sporae fusoideo-ellipsoideae vel fu- 1.5 km n of Pokrent, recultivated sand pit, ~53° 39’ 40’’ n – 11° 8’ 15’’ soideo–oblongatae, oblique monostichae, inaequilaterales vel leniter e, 60 m alt., stem of Melilotus, 13.Xi.2010, leg. t. richter, herb. h.b.

68 Ascomycete.org Fig. 1 – Allophylaria subliciformis (r.M. 2374, photos r. Martínez-Gil). a: habitat of the spanish collection, arrow indicating the exact position where the apothecia were found. b, C: Fresh apothecia, in situ. D: apothecia in the dry state (after 13 months in the herbarium). e–F: Do., margin. G: Medullary excipulum (squash mount). h: ectal excipulum at flanks in external view. all in living state. (e, G, h in water, F in iKi). scale bar = 20 μm for e–h.

Ascomycete.org 69 Fig. 2 Allophylaria subliciformis (h.b. 9456a, photos h.-o. baral). a–D: Fresh apothecia, in situ. e: apothecium in squash mount. F–G: ectal ‒ excipulum in external view, at margin. h: Closeup of marginal cortical cells containing pale yellow, weakly refractive Vbs. i: ectal excipulum in external view, at flanks. J: Do., at junction to stipe. living state (in water), except for F (in Koh). the scale is the same for b–D, F–G, and i–J.

70 Ascomycete.org Fig. 3 Allophylaria subliciformis (r.M. 2374, photos r. Martínez-Gil). a: living mature asci in water. b: Croziers at the ascus base (living state ‒ in water). C–G. Upper part of dead asci in iKi, apices with hemiamyloid apical rings of Laetinaevia-type; C–D: apical rings blue at low iKi concentration; e–G: apical rings red at high iKi concentration. scale bar = 20 μm for all photos.

Ascomycete.org 71 Fig. 4 – Allophylaria subliciformis (a–G, i–l: h.b. 9456a, photos h.-o. baral; h: 12.Xi.2015, photos i. Wagner). a: living mature asci in water. b: Closeup of a. C: Dead mature ascus in iKi at high concentration. D: Do., apices of living mature asci. e: Do., apices of dead immature, mature, and emptied asci, with hemiamyloid (red) apical rings of Laetinaevia-type. F: apices of paraphyses, with subhyaline weakly refractive Vbs. G: living mature ascospores (in iKi). h: Croziers at ascus base (dead state, in Cr). i: living mature ascospores (in water). J: Do., in Crb. K–l: living overmature, 3-septate, germinating ascospores (in water), one spore with chalara-like phialide. the scale is the same for b–l.

72 Ascomycete.org Fig. 5 Allophylaria subliciformis (r.M. 2374, photos r. Martínez-Gil). a: recently expelled spores. b: overmature spores with 1(–3) septa. C: ‒ Germinating spores. D, e: apices of paraphyses containing medium refractive Vbs. F: apices of paraphyses in iKi. all in living state (a–e in water). scale bar = 20 μm for all photos.

9456a. – Thüringen, 2.5 km s of sonneberg, 1 km WsW of oberlind, lipsoid, non-septate spores of 15–26 × 6–7 μm with two large gut- an der Müß, recultivated gravel pit, 50° 20’ 12’’ n – 11° 9’ 55’’ e, 358 m tules. ellis & ellis (1985: 351) reported it on dead stems of Eupato- alt., stem of Solidago canadensis, 12.Xi.2015, leg. i. Wagner (unpre- rium cannabinum l., with fresh apothecia up to 0.7 mm diam. and served, illustrations seen). – Rheinland–Pfalz, 1.7 km nnW of 3-septate spores of 17–29 × 5–7 μm with one large globose oil drop neustadt a.d. Weinstr., haardt, 49° 21’ 47’’ n – 8° 8’ 17’’ e, 195 m alt., in each cell. in spore size and shape these reports are very close to stem of indet. herbaceous dicot, 7.Xii.1975, leg. r. thate, herb. h.b. the free living spores we have documented here. 622. – Baden–Württemberg, 5 km ne of tübingen, 0.6 km s of Dennis (1956: 40) did not observe any iodine reaction of the ascus Pfrondorf, obere Mähder, 48° 32’ 36’’ n – 9° 6’ 40’’ e, 390 m alt., fruit apex in the holotype. the reason is that he used Melzer’s reagent: of Oenothera biennis, 14.i.1990, leg. h.-o. baral (unpreserved, 1 the characteristic hemiamyloid reaction is only visible in iKi but apothecium). – CANADA. Prince-Edward Island, 8 km ssW of Char- masked by the chloral hydrate of Mlz, whereas pretreatment with lottetown, Cumberland, 32 m alt., inflorescence of Cirsium, Koh induces a deep blue reaction that is visible in both iKi and Mlz 30.X.2015, leg. a. Carter, a.C. 2168. (baral, 1987b; s.d./a). bøhler (1974: 81) briefly reported a collection of A. ‘sublicoides’ Comments: Karsten (1869) described the species on the basis of from Aconitum septentrionale and Cicerbita (“Lactuca”) alpina from a single collection on stems of Artemisia vulgaris l., with fusoid-el- the vicinity of oslo, with spores 17–23 × 4–6 µm and inamyloid asci,

Ascomycete.org 73 and an unnamed collection on Chamaenerium angustifolium by possible to distinguish asci which were alive during drying from nannfeldt (Fungi exs. suec. 989) with spores 19–25 × 4.5–6 µm and a those which were dead. deep blue ascus apex. he also compared collections on Athyrium and Dryopteris as “very similar” with those on Aconitum and Cicerbita, Etymology and Nomenclature: in the original description with spores 17–25 × 4–4.5 µm and a faintly blue ascus apex. bøhler (Karsten, 1869, 1870), no reference was made to the meaning of the (op. cit.) used Melzer’s reagent which appears to explain the nega- generic name or to that of the specific epithet. after consulting sev- tive or faintly blue reaction. since he was probably unaware of the eral specialists, we conclude that Allophylaria would mean “different influence of Koh, the observed differences in the iodine reaction and related” in the sense of “originating from a different lineage”, among the studied samples remain doubtful. since he studied only and that the epithet subliciformis derives from the latin feminine herbarium material, the contents of living spores were not taken noun sublica which means a “wooden stake, pile, or pivot”, referring into account in his study. to the similarity in shape of the ascocarps in lateral view with the a recent Canadian collection by a. Carter (pers. comm.) was made supports (timber piles) of the Pons Sublicius (wooden bridge in in october on old inflorescence of Cirsium in a garden. With its asci rome over the tiber river). (†–170 × 12–15 µm) with red iKi reaction and fusoid, slightly curved in the index to his publication in 1869 (p. 205), Karsten briefly sug- spores with rounded apices, *25–30 × 6.5–7.5 µm, 0–3-septate, large gested in a footnote that the specific epithet is to be changed from lbs ~2.5–3 µm diam., this sample appears to fit well (photos by the ‘sublicoides’ (p. 148) to ‘sublicaeformis’. indeed, Karsten (1870) ac- finder can be looked up at baral s.d./b). this seemed us to be the cepted the latter version, and a year later he again changed it to sub- first record of A. subliciformis for north america. Yet, the species al- liciformis (Karsten, 1871). hUhtinen (1990: 242) thought that the index ready appears in a collection list from a foray in indiana, Usa (CooKe, of 1869 was printed somewhat later than the main study, so that 1975) under the misspelled name “Allophylaria subiculoides (Karst.) Karsten could only apply the change as an erratum, but arenDholz nannf.”, identified by r.P. Korf but without indication of the substrate (1989: 289) argued in a footnote that the paper appeared as one en- and without herbarium number. tity and the name change is valid. Despite his doubts, also huhtinen a report by thinD & sinGh (1970) on dead angiospermic wood in argued that the name sublicoides was invalid because it was not ac- himachal Pradesh, india, is too inadequate to recognize its identity: cepted by the author in the original publication (art. 34.1a iCbn st. the apothecia are given up to 0.88 mm diam., the asci as inamyloid, louis = 36.1a iCn Melbourne), so he accepted the variant sublici- and the spores 17–22 × 4–5.4 µm (ascus width is erroneously given formis, as already boUDier (1907) did. also the index Fungorum ac- much too narrow). the lignicolous habitat suggests A. macrospora, cepted this view, because sublicoides was not the original author’s but the euamyloid apical rings of that species contradict this idea. intention and sublicaeformis grammatically incorrect (art. 60.8 iCn a problem with herbarium material is that spore septation cannot Melbourne). nevertheless, various authors (nannFelDt, 1932; Dennis, properly be used as a taxonomic marker, since the percentage of 1956; KorF, 1971; bøhler, 1974; baral & KrieGlsteiner, 1985; ellis & ellis, septate spores strongly depends on the development stage, which 1985; etc.) used the invalid name sublicoides. is difficult to recognise in a dried population. therefore, in some- what senescent apothecia of A. subliciformis most of the asci contain Similar species: rUbio (s.d.) reported Allophylaria byssacea on 1–3-septate spores, whereas apothecia in the optimum stage may stems of Mentha sp. with narrower multiguttulate spores (*16.2–19 hardly show any septate spores. Usually, the living asci contained × 3.7–4.3 μm). he also reported Allophylaria macrospora on wood only non-septate spores in our samples of A. subliciformis. at this of ?Fagus, with spores similar in size to A. sublicoides (*25.5–28.3 × stage, which was termed mature by baral (1992), they are ejected. 5.6–6.6 μm) but multiguttulate and with asci with a simple-septate however, since many asci die under natural condition or in a moist base (for these two species see also baral s.d./b). rubio’s statement box before they succeed to eject their spores, many still living spores of blue apical rings in iKi of high concentration but red in iKi of low may get overmature within the dead asci after some time. they are concentration is surely a confusion, since the hemiamalyoid reaction still there when studying a herbarium specimen, in which it is im- of type rb is always the other way round: blue at low and red at high

Table 1 – Comparison of selected characters in the five european collections

Spain Mecklenburg Tübingen Rheinland-Pfalz Thüringen *(145–)150–171(–175) *157–205 × 15.5–16.5 *180–240 × 16–17.8 *158–177 × 16–19 µm Asci × (15–)15.5–16.5(–17) µm, †135-150 × (10–) *190 × 15–17.5 µm µm †150 × 11–15 µm µm 11.5-13.5 µm IKI reaction ~rb (strong) rb (strong) ~rr (strong) rb (strong) ~rb (strong) *(22–)24–26(–28) × *23.5–25(–27.5) × 6– *(19.5–)21–26(–28) × *20–25(–26.5) × 6– Spores 6.5–7(–7.5) µm, †(17–) *20.5–25 × 6.3–7 µm 7 µm, †19–23 × 5– (6–)6.5–7(–7.5) µm 8 µm 18–23(–24 × 5–6.3 µm 6 µm Sheath + + + ? + 2.5–4 4–5.5 µm 3–4 µm 3–4.5 µm ~2–4 µm Large LBs → 2.8–4.3 µm 1–3 per half 1 per half 1(–2) per half ?1 per half

Septa 0 → 1–3 0–1 → 1–3 0 → 1(–3) → 1–3 0 → 1–3 Paraph. apex *4.2–5.7 μm *3–5 μm *3–5 μm ? *2–4 μm width 17–30 µm long 10–20 µm long 17–30 µm long 15–25 µm long Paraph. VBs hyaline, iKi yellow- iKi red-brown guttu- ? pale chlorinaceous hyaline orange guttulate late (0.15–)0.25–0.35 Apoth. diam. 0.2–0.5 mm (0.25)0.3–0.4(–0.5) mm ? 0.2–0.3 mm (–0.4) mm Host indet. dicot Melilotus Oenothera biennis indet. dicot Solidago canadensis

74 Ascomycete.org concentration. A. macrospora was found by baral (s.d./b) to deviate deutschland gefundene inoperculate Discomyzeten, mit tax- from A. subliciformis by reacting bb, i.e., blue in a wide range of con- onomischen, ökologischen, chorologischen hinweisen und einer centration by turning dirty reddish only above ~0.5% or not even Farbtafel. Beihefte zur Zeitschrift für Mykologie, 6: 1–160. then, whereas A. subliciformis changes from blue to red already baral h.-o. s.d./a. — iodine reaction in ascomycetes: why is lugol’s around 0.2–0.3%. solution superior to Melzer’s reagent? https://invivoveritas.de/articles/io- dine–reaction–in–ascomycetes–why–is–lugols–solution–superior–to–melzers– Ecology: Allophylaria subliciformis appears to prefer open ruderal reagent/ vegetation, where it grows on previous year’s broken stems of baral h.-o. s.d./b. — In vivo veritas. over 10.000 scans of fungi and herbaceous dicots lying on the periodically moist ground in pas- plants, with materials on vital taxonomy and desiccation toler- tures. two collection sites were large pits in which sand or gravel ance. 3rd ed., DVD-roM. also available on https://invivoveritas.de was exploited and which got naturally recultivated. the samples bøhler h.C. 1974. — taxonomical studies on some norwegian Helo- listed here were all from late autumn and winter, and the different tiales (ascomycetes) on fern remains. Norwegian Journal of Botany, hosts indicate that the species is not substrate-specific but grows 21: 79–100. on hosts of different families (Fabaceae, Oenotheraceae, Asteraceae), boUDier É. 1907. — Histoire et classification des discomycètes d’Europe. all being typical ruderal plants. the type was collected in the end of Paris, P. Klincksieck, 223 p. october on desiccated stems of Artemisia (Karsten, 1869). Under the CarPenter s.e. 1981. — Monograph of Crocicreas (ascomycetes, Helo- name Cyathicula subliciformis the Fungal records Database of britain tiales, Leotiaceae). Memoirs of the New York Botanical Garden, 33: and ireland lists 11 records belonging to about 8 samples, partly de- 1–290. posited in K(M), collected in september and october on Eupatorium CooKe W.b. 1975. — the 1970 indian Foray. Mycologia, 67 (5): 1065– and unidentified herbs. the correctness of these records as those 1071. referred by bøhler (1975) to this species remains to be clarified. Fur- Dennis r.W.G. 1956. — a revision of the british Helotiaceae in the ther records did not come to our attention. herbarium of the royal botanic Gardens, Kew, with notes on re- lated european species. Mycological Papers, 62: 1–216. Acknowledgements ellis M.b. & ellis J.P. 1985. — Microfungi on land plants. an identifi- cation handbook. london & sydney, Croom helm, 818 p. Grelet l.-J. 1951. — les Discomycètes de France d’après la classifi- Guy Marson is thanked for gaining an rDna sequence, Martin be- cation de boudier (22e fascicule). Revue de Mycologie, 16: 80–100. mmann for literature research and proofreading the manuscript, hUhtinen s. 1990. — a monograph of Hyaloscypha and allied genera. and adrian Carter, torsten richter, ingo Wagner, and the late rudolf Karstenia, 29: 45–252. thate for supplying their collections and observations. We are grate- JaKlitsCh W., baral h.-o., lüCKinG r. & lUMbsCh h.t. 2016. — ascomy- ful to Paul Kirk, shaun Pennycook, Chris Yeates, and Viktorie halasů cota. In: FreY W. (ed.). Engler’s syllabus of plant families. 13th ed., Part for their help in understanding the etymology of subliciformis, and 1/2. stuttgart, borntraeger, 322 p. to Fernando Martínez for searching and translating some docu- Karsten P.a. 1869. — Monographia Pezizarum fennicarum. Notiser ur ments. Sällskapets pro Fauna et Flora fennica förhandlingar, 10: 99–206. Karsten P.a. 1870. — symbolae ad mycologiam Fennicam i. Notiser References ur Sällskapets pro Fauna et Flora Fennica Förhandlingar, 11: 211– 268. arenDholz W.-r. 1989. — the genus Pezizella i: nomenclature and Karsten P.a. 1871. — Mycologia fennica i. Discomycetes. Bidrag till history. Mycotaxon, 36: 283–303. kännedom af Finlands natur och folk, 19: 1–263. baral h.-o. 1987a. — Der apikalapparat der Helotiales. eine licht- Karsten P.a. 1885. — revisio monographica atque synopsis as- mikroskopische studie über arten mit amyloidring. Zeitschrift für comycetum in Fennia hucusque detectorum. Acta Societas Pro Mykologie, 53 (1): 19–135. Fauna et Flora Fennica, 2: 1–174. baral h.-o. 1987b. — lugol’s solution/iKi versus Melzer’s reagent: KorF r.P. 1971. — some new discomycete names. Phytologia, 21 (4): hemiamyloidity, a universal feature of the ascus wall. Mycotaxon, 201–207. 29: 399–450. nannFelDt J.a. 1932. — studien über die Morphologie und system- baral h.-o. 1992. — Vital versus herbarium taxonomy: Morpholog- atik der nichtlichenisierten inoperculaten Discomyceten. Nova ical differences between living and dead cells of ascomycetes, and Acta Regiae Societatis Scientiarum Upsaliensis, ser. IV, 8 (2): 1–368. their taxonomic implications. Mycotaxon, 44: 333–390. rabenhorst l. 1869. — repertorium. P.a. Karsten, Monographia baral h.-o. 2015. — nomenclatural novelties: new combinations in Pezizarum fennicarum. Hedwigia, 8 (11): 162–166. the Leotiomycetes. Index Fungorum, 271: 1. rUbio e. s.d. — Allophylaria. asturnatura. https://www.asturnatura.com/gen- baral h.-o. & räMä t. 2015. — Morphological update on Calycina ma- ero/allophylaria.html [accessed 4 oct. 2017]. rina (Pezizellaceae, Helotiales, Leotiomycetes), a new combination thinD K.s. & sinGh h. 1970. — the Helotiales of india XiV. Proceedings for Laetinaevia marina. Botanica Marina, 58 (6): 523–534. doi: of the Indian Academy of Science, sect. B, 72: 171–178 + Pl. Viii. 10.1515/bot-2015-0049 triebel D. & baral h.-o. 1996. — notes on the ascus types in Croci- baral h.-o. & KrieGlsteiner G.J. 1985. — bausteine zu einer creas (Leotiales, ascomycetes) with a characterization of selected askomyzeten–Flora der bundesrepublik Deutschland: in süd- taxa. Sendtnera, 3: 199–218. ef

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1: R. Martínez-Gil – Parque San Miguel Nº12, 2ºA, La Rioja, 26007 Lagroño, Spain – [email protected] 2: H.-O. Baral – Blaihofstr. 42, 72074 Tübingen, Germany – [email protected]

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