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Phaeocalicium (Mycocaliciaceae, Ascomycetes) in Northern Europe

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Phaeocalicium (Mycocaliciaceae, Ascomycetes) in Northern Europe Ann. Bot. Fennici 33: 205–221 ISSN 0003-3847 Helsinki 30 October 1996 © Finnish Zoological and Botanical Publishing Board 1996 Phaeocalicium (Mycocaliciaceae, Ascomycetes) in Northern Europe Leif Tibell Tibell, L., Department of Systematic Botany, Villavägen 6, S-752 36 Uppsala, Sweden Received 7 February 1996, accepted 28 May 1996 The taxonomy, distribution and ecology of eight species of Phaeocalicium A. F. W. Schmidt (Mycocaliciaceae, Ascomycetes) occurring in the Nordic countries and Greenland are de- scribed. They are parasitic or saprophytic mainly on thin twigs of trees and shrubs such as Alnus, Betula, Populus and Salix and are often quite host-specific. A key to the species is supplied. Two new species, Ph. boreale Tibell and Ph. flabelliforme Tibell are described. A lectotype is selected for Calicium praecedens Nyl., Mycocalicium pusiolum (Ach.) Räsänen var. macrospora Räsänen and Stenocybe tremulicola Norrl. ex Nyl., and a neotype is se- lected for Phaeocalicium populneum (Brond. ex Duby) A. F. W. Schmidt. The new combi- nation Ph. tremulicola (Norrl. ex Nyl.) Tibell is proposed. Key words: Ascomycetes, Caliciales, Mycocaliciaceae, Phaeocalicium, taxonomy INTRODUCTION cybe, as conceived by Schmidt (1970), are unsatis- factory and in need of revision. Thus as an example The ascomycete genus Phaeocalicium belongs to spore septation does not seem to be consistent evi- Mycocaliciaceae in Caliciales s. lat. (Tibell 1984). dence for generic delimitation. In the present paper The North European species occur as saprobes and/ a group of similar and presumably closely related or weak parasites mostly on thin, decaying branches species has provisionally been accomodated in of deciduous trees or shrubs. The genus occurs Phaeocalicium pending a detailed analysis based on mainly in cool temperate to temperate areas of the macromolecular evidence in progress. Notwithstand- Northern Hemisphere, with one species each occur- ing a regional revision of Phaeocalicium in the USSR ring in Australasia and South America. Phaeo- by Titov (1986), the genus is still incompletely calicium was described by Schmidt (1970) and in known. In this paper eight species occurring in North- that paper Ph. compressulum, Ph. populneum and ern Europe are treated, two of which are previously Ph. praecedens were dealt with in detail. As con- undescribed. ceived there Phaeocalicium includes species in Mycocaliciaceae with rather large, simple or 1- septate spores, large asci and a strongly and evenly MATERIAL AND METHODS thickened ascus apex. As noted earlier (Tibell 1984, 1995), the generic relationships between Phaeoca- The study has been based on field studies and material col- licium, Chaenothecopsis, Mycocalicium and Steno- lected by the author (kept in UPS), additional material from 206 Tibell • ANN. BOT. FENNICI 33 (1996) the herbaria at C, H, LD, O, S, UPS, as well as the private — Excipulum consisting of periclinally arranged hyphae, herbaria of S. Hultengren (Stenungsund), A. Nordin (Uppsala), without isodiametric cells, spores dark brown, with a M. Karström (Vuollerim), L.-E. Muhr (Karlskoga), and D.- minute ornamentation .................. 3. Ph. compressulum O. Övstedal (Bergen). 4. Excipulum edge not thickened, ascomata 0.6–0.9 mm Estimates of the size of ascomata height, capitulum high, on Populus ............................ 7. Ph. praecedens width, stalk width, ascus length, ascus width, spore length — Excipulum edge strongly thickened, ascomata 0.3–0.4 mm (l) and spore width (w) are given as l––l+ × w––w+ where l– high, on Populus and Salix ................. 5. Ph. interruptum and w– represent the arithmetic mean minus 1 standard de- 5. Capitula strongly flattened ......... 4. Ph. flabelliforme viation and l+ and w+ the arithmetic mean plus 1 standard — Capitula not strongly flattened..................................6 deviation. If the measurements have been estimated by sta- 6. Ascomata 0.4–0.7 mm high, excipulum consisting of tistical methods they are followed by a parenthesis in which periclinally arranged hyphae, on Populus .................. the arithmetic mean (X), the standard deviation (s), the num- ....................................................... 6. Ph. populneum ber of observations (n) and the number of collections meas- — Ascomata 0.3–0.4 mm high, excipulum consisting of ured (co) are given. If statistical estimates are not given the isodiametric or periclinally arranged cells, on Alnus, sizes indicated represent common ranges. Betula, Populus and Salix .........................................7 7. Excipulum edge strongly thickened, spores 9–11 × 4– 4.5 µm ........................................... 5. Ph. interruptum — Excipulum edge not strongly thickened, spores 11–17 PHAEOCALICIUM A. F. W. SCHMIDT × 4–6 µm ...................................................................8 8. Excipulum and stalk reddish in section, K+ intensified Mitt. Staatsinst. Allg. Bot. Hamburg 13: 128. 1970. — Type: reddish, stalk finally dark reddish grey ....... 2. Ph. boreale Calicium praecedens Nyl. = Phaeocalicium praecedens — Excipulum and stalk pale, brownish or greenish brown (Nyl.) A. F. W. Schmidt. in section, K– ............................................................9 9. Mature spores with heavily pigmented septa, excipulum Saprophytic or parasitic. Ascomata well stalked, consisting of a single layer of isodiametric cells with thick black to blackish brown. Capitulum obovoid to len- walls 5–6 µm wide...........................8. Ph. tremulicola ticular or strongly compressed. Excipulum well de- — Mature spores with poorly pigmented septa, excipulum veloped, consisting of dark brown, periclinally ar- consisting of 2–3 layers of sclerotized, periclinally ranged or isodiametric hyphae. Hymenium covered arranged hyphae......................... (Stenocybe pullatula) by a thin epithecium. Stalk consisting of dark brown to aeruginose or pale, periclinally arranged hyphae. 1. Phaeocalicium betulinum (Nyl.) Tibell (Fig. 1) Asci narrowly cylindrical, with uniseriately arranged spores, 70–100 µm long, formed singly from asco- Ann. Bot. Fennici 28: 118. 1991. — Calicium pallescens genous hyphae with hooks. Asci with strongly and Nyl. f. betulinum Nyl., Flora 50: 370. 1867. — Type: Fin- uniformly thickened apex or the apex is penetrated land. Tavastia australis, Asikkala, 1866 Norrlin (H-NYL by a short and blunt canal persisting until the spores 40730, lectotype, designated by Tibell, Ann. Bot. Fennici are ejected. No mazaedium. Spores 10–18 × 4–6 µm, 28: 118. 1991; isotypes: H). non-septate or 1–3-septate, ellipsoidal, with rounded Saprophytic or parasitic on Betula. Ascomata 0.27– apices or fusiform. Spore wall rather thick, dark 0.36 mm high (X = 0.31 mm, s = 0.05 mm, n = 20, brown, smooth or with a verrucose ornamentation, co = 3), shiny. Capitulum blackish brown, strongly flat- not rupturing at maturity. Conidiomata unknown. tened, 0.08–0.13 mm wide (X = 0.11 mm, s = 0.03 mm, Chemistry. No secondary substances identi- n = 20, co = 3), epruinose. Epithecium brown, 6–8 µm fied. Some species contain pigments which change thick. Hypothecium c. 75 µm high, hyaline, consist- colour with a change of pH yielding reactions with ing of mainly periclinally arranged, winding and KOH and HNO3 observable under the microscope. sparsely branched, thin-walled hyphae, 1.5–2 µm diam. Excipulum medium brown, 14–25 µm thick, composed of two layers. The outer layer is formed Key to the species by isodiametric to irregular, slightly sclerotized cells with strongly thickened walls and lumina only 1–2 µm 1. Spores non-septate ....................................................2 diam. The inner part of the excipulum consists of 2–4 — Spores 1(–3)-septate .................................................5 layers of periclinally arranged hyphae 1–1.5 µm 2. Capitula strongly flattened ........................................3 — Capitula lenticular .................................................... 4 diam. Excipulum and epithecium HNO3– and K–. 3. Excipulum with an outer layer of small, isodiametric Stalk pale, olivaceous brown, 0.04–0.05 mm diam., cells, spores pale brown, smooth ..... 1. Ph. betulinum K–, HNO3–, in section very pale olivaceous brown, ANN. BOT. FENNICI 33 (1996) • Phaeocalicium in Northern Europe 207 Fig. 1. Phaeocalicium betulinum (Nyl.) Tibell. — A: Longitudinal section of ascoma. — B: Longitudinal section of ascoma stalk with gelatinous coat. Stalk moderately pigmented, consisting of periclinally arranged, short-celled, slightly intertwined hyphae. — C: Excipulum consisting of two layers. The outer layer (o) is formed by isodiametric to irregular, slightly sclerotized, thick-walled cells, the inner part (i) by 2–4 layers of periclinally arranged, sclerotized hyphae. — D, E: Mature asci. Phase contrast micrograph. — F–H: Mature spores. Under the light microscope the spores appear smooth. In TEM the almost mature spores have a wall consisting of three layers: an outermost, thin layer with a very electron-dense lining (L); a thick middle layer with irregularly distributed pigment granules (M); and an electron-lucent innermost layer (i). Surface of the spore only very slightly and irregularly uneven. In SEM the spore surface with a very minute and irregular ornamentation partly consisting of longitudinally arranged wrinkles. — A–D, F: Lectotype; E, G, H: isotype, H. — Scales: A: 50 µm; B: 25 µm; C–F: 20 µm; G–H: 2 µm. consisting of periclinally arranged, short-celled, Asci 47.6–70.7 × 4.4–5.1 µm (X = 59.2 µm, slightly intertwined hyphae,
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