Jaklitsch WM and Voglmayr H (2014), Persistent hamathecial threads in the Nectriace...

ECB-ART-43862

Persoonia 2014 Dec 01;33:182-211. doi: 10.3767/003158514X685211.

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Persistent hamathecial threads in the Nectriaceae, Hypocreales: Thyronectria revisited and re-instated.

Jaklitsch WM , Voglmayr H .

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Based on type studies and freshly collected material we here re-instate the genus Thyronectria (Nectriaceae, Hypocreales). Species of this genus were recently for the most part classified in the genera Pleonectria (Nectriaceae) or Mattirolia (Thyridiaceae), because Thyronectria and other genera had been identified as members of the Thyridiaceae due to the presence of paraphyses. Molecular phylogenies based on several markers (act, ITS, LSU rDNA, rpb1, rpb2, tef1, tub) revealed that the Nectriaceae contain members whose ascomata are characterised by long, more or less persistent, apical paraphyses. All of these belong to a single genus, Thyronectria, which thus has representatives with hyaline, rosy, green or even dark brown and sometimes distoseptate ascospores. The type species of Thyronectria, T. rhodochlora, syn. T. patavina, syn. T. pyrrhochlora is re-described and illustrated. Within the Nectriaceae persistent, apical paraphyses are common in Thyronectria and rarely also occur in Nectria. The genus Mattirolia is revised and merged with Thyronectria and also Thyronectroidea is regarded as a synonym of Thyronectria. The three new species T. asturiensis, T. caudata and T. obscura are added to the genus. Species recently described in Pleonectria as well as some species of Mattirolia are combined in the genus, and a key to Thyronectria is provided. Five species are epitypified. The type species of the genus Thyridium (Thyridiaceae), T. vestitum, is included in phylogenetic analyses to illustrate the phylogenetic distance of Thyronectria from the Thyridiaceae.

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Species referenced: Echinodermata
Genes referenced: LOC100888242 LOC583794 LOC589952 LOC590297

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	Fig. 1. Phylogram showing one of four MP trees 9 282 steps long revealed by PAUP from an analysis of the combined 4-gene (LSU, rpb1, rpb2, tef1) matrix of selected Xylariomycetidae, Sordariomycetidae and Nectriacae, showing the phylogenetic position of Thyridium, Thyronectria and Mattirolia (given as Thyronectria roseovirens in the tree). MP and ML bootstrap support above 50 % and Bayesian posterior probabilities above 90 % are given above or below the branches. Strain/culture designations are given following the taxon names; new species are marked in bold italics. Nodes marked by an asterisk (*) collapsed in the strict consensus of the four MP trees.
	Fig. 2. Phylogram showing one of 216 MP trees 6 654 steps long revealed by PAUP from an analysis of the combined 7-gene (act, ITS, LSU, rpb1, rpb2, tef1, tub) matrix of Allantonectria and Thyronectria, with four species of Nectria s.str. selected as outgroup. MP and ML bootstrap support above 50 % and Bayesian posterior probabilities above 90 % are given above or below the branches. Strain/culture designations are given following the taxon names; new species are formatted in bold; holo-, neo- or epitype strains/specimens are marked by an asterisk (*).
	Fig. 3. Thyronectria rhodochlora, stromata and ascomata. a–h, j–m. Dry stromata in surface view (a. habit; b. with pycnidia of a Diplodia sp.); i. stromata in 3 % KOH after rehydration; n. perithecium in vertical section; o. lower lateral portion of a perithecium showing asci and apical paraphyses in section; p–r. peridium in section (q. basal region; r. ostiolar region with periphyses and scurf); s. stroma hyphae (n–p, r: in 50 % glycerol; q, s: in lactic acid). a, e, i, n–s: epitype WU 31656 (NP3); b, c, g: WU 32152 (NP7); d, f, h: WU 31654 (NP1); j, k: lectotype (PC); l: isolectotype (K); m: holotype of T. patavina (PAD). — Scale bars: a, m = 1 mm; b, e, h, i = 0.4 mm; c, d, f, j–l = 0.3 mm; g = 0.7 mm; n = 100 μm; o, q = 30 μm; p, r, s = 15 μm.
	Fig. 4. Thyronectria rhodochlora, centrum characteristics. a–d. Apical paraphyses among immature (a, b) and mature (c, d) asci in water; e–j. asci with ascospores; k–s. ascospores (q. overmature, producing conidia; all in water except p (in water after swelling in 3 % KOH) and s. (in lactic acid in a microtome section)). a, e, f, k, l: T. patavina holotype (PAD); b: WU 32150 (NP5); c, h, i, o, s: epitype WU 31656 (NP3); d, p: T. rhodochlora isolectotype (K); g, n, r: WU 31654 (NP1); j: WU 32152 (NP7); m: T. rhodochlora lectotype (PC)); q. WU 33425a (NP8). — Scale bars: a, e, h = 20 μm; b = 30 μm; c = 50 μm; d, f, g, i, j, m, p = 15 μm; k, n, o = 10 μm; l, q–s = 7 μm.
	Fig. 5. a–l: Thyronectria rhodochlora, asexual morph. a, b. Pycnidia (MEA, 15 °C, 65 d); c–h. conidiophores and phialides (c, d, h. from pycnidium on MEA, 15 °C, 65 d; c. shrub-like; e–g. from effuse conidiation, showing pegs and phialides; MEA, RT, 3 d); i–l. conidia (i, j: MEA, RT, 3 d; k, l: from pycnidium on MEA, 15 °C, 65 d) (all in water except h (in 3 % KOH)). a–d, h, k, l: NP7; e–g, i, j: NP. — m–s. Thyronectria virens (isolectotype BPI 631193). m–o. Ascomata in bark (o. old open ascoma attached to the side of a Diplodia conidioma); p–s. ascospores. — t–w. Ascospores of Thyronectria cf. virens (WU 33426; u. in ascus, note apical paraphyses). — x–aa. Thyronectria zanthoxyli. x, y. Stromata; z, aa. ascospores. x, y, aa. lectotype NYS 3611; z. isolectotype NYS 3438. — Scale bars: a, m, n, x = 0.5 mm; b, o, y = 0.3 mm; c = 20 μm; d, f = 15 μm; e, g–j, p, q, t, u, v, z = 10 μm; k, l, r, s, w, aa = 7 μm.
	Fig. 6. Thyronectria asturiensis WU 32124 (MA3). a–f. Stromata/ascomata (f. in 3 % KOH); g. perithecium in vertical section; h–k. asci with ascospores; l–n. peridium in section (l. ostiolar region; m. base; n. lateral; l. in lactic acid; m, n. in 3 % KOH); o. ascospores; p, q. conidiophores and phialides (20 °C, MEA, 6 d); r. conidia (20 °C, MEA, 6 d). — Scale bars: a, b, e = 0.3 mm; c, d = 0.2 mm; f = 0.5 mm; g = 100 μm; h, i, k, o, q = 10 μm; j, l, p = 15 μm; m, n = 20 μm; r = 5 μm.
	Fig. 7. a–d: Thyronectria austroamericana WU 32664. a, b. Pycnidial aggregates on hypostromata; c. phialides; d. conidia. — e–p: Thyronectria chrysogramma. e–h. Stromata (e, f. holotype; g, h. paratype); i, n–p. ascospores; j. apical paraphyses and asci; k–m. asci (k, l. immature; k. showing oblong ascospores). — q–w. Thyronectria coryli. q, r. Perithecia with greenish yellow scurf; s. apical paraphyses and mature asci; t. ascospores and ascoconidia within asci; u. ascoconidia; v, w. asci with ascospores and ascoconidia (q: WU 32129; r: WU 30362 (NCP); s–w: WU 32127 (W.J. 1262)). — Scale bars: a, b = 1.5 mm; c, i, w = 10 μm; d, u = 5 μm; e–g, q = 0.4 mm; h, r = 0.2 mm; j = 30 μm; k–p, t, v = 15 μm; s = 75 μm.
	Fig. 8. Thyronectria caudata, sexual morph. a–f. Perithecial aggregates (f. in 3 % KOH); g, l. perithecia and hypostroma in vertical section; h, i. peridium in section (3 % KOH; h. ostiolar region); j. hypostroma cells (3 % KOH); k. host hyphae (lactic acid); m. mature asci and apical paraphyses; n, o. asci with ascospores and ascoconidia; p, q. ascospores; r. ascoconidia. a, b, d–m, o, q, r: epitype WU 32130; c, p: holotype (MPU); n. BPI 552469. — Scale bars: a = 0.5 mm; b–f = 0.3 mm; g = 0.2 mm; h, m = 30 μm; i–k, o = 20 μm; l = 100 μm; n = 15 μm; p, q = 10 μm; r = 5 μm.
	Fig. 9. Thyronectria caudata WU 32130 (NL2), asexual morph. a, b. Pycnidia on the natural substrate (b. in association with perithecia); c. pycnidia on CMD; d–l. conidiophores and phialides (d, g. from natural host; e, f, h–l. from MEA, 20 °C, 7–9 d); m–p. conidia (m, o. from MEA, 20 °C, 7 d; n, p. from the natural host). — Scale bars: a, c = 0.2 mm; b = 0.3 mm; d, h, i, l, n = 7 μm; e, k = 15 μm; f, j = 10 μm; g, m, o, p = 5 μm.
	Fig. 10. a–n: Thyronectria lamyi. a–f. Perithecial aggregates (a. immature; d. with an ascoma of Cucurbitaria berberidis at the upper end; e. erect pycnidium surrounded by perithecia; f. superficial perithecia after bark removal); g, h. asci with ascospores and ascoconidia; i. apical paraphyses at ascus bases; j. ascoconidia; k, l. budding ascospores; m, n. sterile hyphae and conidia from pycnidium on natural host (a: epitype WU 32159 (NL3); b–n: WU 32141 (NL1)). — o, p: Mattirolia maclurae (holotype, LISE). o. ascus; p. ostioles in surface view. — Scale bars: a, c, e, f = 0.25 mm; b, d = 0.4 mm; g–i, m, o = 15 μm; j, n = 5 μm; k, l, p = 10 μm.
	Fig. 11. Thyronectria obscura. a–h. Stromata/Perithecia (h. in 3 % KOH); i. perithecium in vertical section; j, k. peridium in section in 3 % KOH (j. ostiolar region; k. base); l. stroma hyphae in lactic acid; m–o. asci with ascospores and ascoconidia, and apical paraphyses; p, q. ascospores; r–t. conidiophores and phialides (MEA, RT, 2d); u. ascoconidia; v, w. conidia (MEA, RT, 2 d) (a–e, h–l, n, p, r–t, v, w: WU 32142 (TT); f, g, m, o, q, u: WU 32143 (TT1)). — Scale bars: a, c, d, f = 0.2 mm; b, e, g, h = 0.3 mm; i = 50 μm; j–l = 25 μm; m–o = 15 μm; p–s, v, w = 10 μm; t, u = 5 μm.
	Fig. 12. Thyronectria roseovirens, ascomata and stromata (h, i, k. showing rosy asexual morph). Note direct growth on the host fungus Cucurbitaria laburni in a (a–d, o: WU 32154 (MA1; o. in 3 % KOH); e, k: WU 32153 (MA); f, g: lectotype (FH); h, i, m: WU 32156 (MA4); j: WU 32157 (MA5); l: WU 32155 (MA2); n: WU 32158 (MA6)). — Scale bars: a, d, g = 0.3 mm; b, c, e, f, h–o = 0.5 mm.
	Fig. 13. Thyronectria roseovirens, microscopic traits. a. Section of perithecium including natural host; b. perithecium in vertical section; c–e. peridium in section (c. ostiolar region with periphyses in water; d. lateral region in lactic acid; e. base in water); f. stroma cells in lactic acid; g–k. ascospores (g. immature, with projecting sheath; h. showing distosepta); l. apical paraphyses; m–s. asci, ascospores and apical paraphyses; t–w. conidiophores and phialides (MEA, RT, 3–4 d); x, y. conidia (MEA, RT, 3–4 d) (a–f, h: WU 32154 (MA1); g, i: WU 32158 (MA6); j, o, p, s: WU 32155 (MA2); k–n: lectotype (FH); q: WU 32156 (MA4); r: WU 32157 (MA5); t–y: WU 32153 (MA)). — Scale bars: a = 100 μm; b = 50 μm; c, e, p–s, v = 20 μm; d, h, l–o = 15 μm; f, i–k, t, x, y = 10 μm; g, u, w = 5 μm.

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