Research Article |
Corresponding author: Walter M. Jaklitsch ( walter.jaklitsch@univie.ac.at ) Academic editor: Huzefa Raja
© 2019 Walter M. Jaklitsch, Hermann Voglmayr.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Jaklitsch WM, Voglmayr H (2019) European species of Dendrostoma (Diaporthales). MycoKeys 59: 1-26. https://doi.org/10.3897/mycokeys.59.37966
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European species of the genus Dendrostoma (Erythrogloeaceae, Diaporthales) occurring on Castanea sativa and Quercus spp. based on freshly collected material are presented. Using a matrix of sequences from ITS, LSU, rpb2, and tef1, five species are recognized, and their phylogenetic positions are determined. Four species are added to the 14 described species of Dendrostoma. Dendrostoma atlanticum on Castanea sativa, D. creticum on Quercus coccifera and D. istriacum on Q. ilex are described as new species, Valsa castanea is combined in Dendrostoma, and D. leiphaemia is redescribed and illustrated. A key to the European species of Dendrostoma is provided.
Amphiporthe, Cryptodiaporthe, multi-gene phylogeny, pyrenomycetes, systematics
The genus Cryptodiaporthe, based on Cryptospora aesculi, is one of several segregates from the large genus Diaporthe (Diaporthales), characterized by the lack of stromatic zones and with asexual morphs recognized by
All isolates included in this study originated from ascospores of freshly collected specimens derived from recently dead branches or twigs. Details of the strains including NCBI GenBank accession numbers of gene sequences used to compute the phylogenetic trees are listed in Table
Isolates and accession numbers of sequences used in the phylogenetic analyses.
Species | Culture1,2 | Country | Host | Host family | GenBank accession numbers2 | |||
---|---|---|---|---|---|---|---|---|
ITS | LSU | rpb2 | tef1 | |||||
Chrysocrypta corymbiae | CBS 132528* | Australia | Corymbia sp. | Myrtaceae | JX069867 | JX069851 | MH545415 | MH545457 |
Dendrostoma atlanticum | D196 = CBS 145804* | France | Castanea sativa | Fagaceae | MN447223 | MN447223 | MN432160 | MN432167 |
D303 | Spain | Castanea sativa | Fagaceae | MN447224 | MN447224 | MN432161 | MN432168 | |
Dendrostoma aurorae | CFCC 52753* | China | Castanea mollissima | Fagaceae | MH542498 | MH542646 | MH545405 | MH545447 |
CFCC 52754 | China | Castanea mollissima | Fagaceae | MH542499 | MH542647 | MH545406 | MH545448 | |
Dendrostoma castaneae | CFCC 52745* | China | Castanea mollissima | Fagaceae | MH542488 | MH542644 | MH545395 | MH545437 |
CFCC 52746 | China | Castanea mollissima | Fagaceae | MH542489 | – | MH545396 | MH545438 | |
CFCC 52747 | China | Castanea mollissima | Fagaceae | MH542490 | – | MH545397 | MH545439 | |
CFCC 52748 | China | Castanea mollissima | Fagaceae | MH542491 | – | MH545398 | MH545440 | |
CFCC 52749 | China | Castanea mollissima | Fagaceae | MH542492 | MH542645 | MH545399 | MH545441 | |
CFCC 52750 | China | Castanea mollissima | Fagaceae | MH542493 | – | MH545400 | MH545442 | |
CFCC 52751 | China | Castanea mollissima | Fagaceae | MH542494 | – | MH545401 | MH545443 | |
CFCC 52752 | China | Castanea mollissima | Fagaceae | MH542495 | – | MH545402 | MH545444 | |
Dendrostoma castaneicola | CFCC 52743* | China | Castanea mollissima | Fagaceae | MH542496 | – | MH545403 | MH545445 |
CFCC 52744 | China | Castanea mollissima | Fagaceae | MH542497 | – | MH545404 | MH545446 | |
Dendrostoma castaneum | D192 = CBS 145803 | Austria | Castanea sativa | Fagaceae | MN447225 | MN447225 | MN432162 | MN432169 |
D230 | Italy | Castanea sativa | Fagaceae | MN447226 | MN447226 | – | MN432170 | |
D260 | Italy | Castanea sativa | Fagaceae | MN447227 | MN447227 | – | – | |
Dendrostoma chinense | CFCC 52755* | China | Castanea mollissima | Fagaceae | MH542500 | MH542648 | MH545407 | MH545449 |
CFCC 52756 | China | Castanea mollissima | Fagaceae | MH542501 | MH542649 | MH545408 | MH545450 | |
CFCC 52757 | China | Castanea mollissima | Fagaceae | MH542502 | MH542650 | MH545409 | MH545451 | |
CFCC 52758 | China | Castanea mollissima | Fagaceae | MH542503 | MH542651 | MH545410 | MH545452 | |
Dendrostoma creticum | D124 = CBS 145802* | Greece | Quercus coccifera | Fagaceae | MN447228 | MN447228 | MN432163 | MN432171 |
Dendrostoma dispersum | CFCC 52730* | China | Quercus sp. | Fagaceae | MH542467 | MH542629 | MH545374 | MH545416 |
CFCC 52731 | China | Quercus sp. | Fagaceae | MH542468 | MH542630 | MH545375 | MH545417 | |
Dendrostoma istriacum | D122 = CBS 145801* | Croatia | Quercus ilex | Fagaceae | MN447229 | MN447229 | MN432164 | MN432172 |
Dendrostoma leiphaemia | D105 = CBS 145800 | Austria | Quercus robur | Fagaceae | MN447230 | MN447230 | MN432165 | MN432173 |
D144 | Poland | Quercus robur | Fagaceae | MN447231 | MN447231 | MN432166 | MN432174 | |
CBS 187.37 | NA | Quercus sp. | Fagaceae | MH855882 | MH867393 | – | – | |
Dendrostoma mali | CFCC 52102* | China | Malus spectabilis | Rosaceae | MG682072 | MG682012 | MG682032 | MG682052 |
Dendrostoma osmanthi | CFCC 52106* | China | Osmanthus fragrans | Oleaceae | MG682073 | MG682013 | MG682033 | MG682053 |
CFCC 52107 | China | Osmanthus fragrans | Oleaceae | MG682075 | MG682015 | MG682035 | MG682055 | |
CFCC 52108 | China | Osmanthus fragrans | Oleaceae | MG682074 | MG682014 | MG682034 | MG682054 | |
CFCC 52109 | China | Osmanthus fragrans | Oleaceae | MG682076 | MG682016 | MG682036 | MG682056 | |
Dendrostoma parasiticum | CFCC 52761 | China | Castanea mollissima | Fagaceae | MH542480 | MH542636 | MH545387 | MH545429 |
CFCC 52762* | China | Quercus wutaishanica | Fagaceae | MH542482 | MH542638 | MH545389 | MH545431 | |
CFCC 52763 | China | Castanea mollissima | Fagaceae | MH542481 | MH542637 | MH545388 | MH545430 | |
CFCC 52764 | China | Quercus aliena | Fagaceae | MH542483 | MH542639 | MH545390 | MH545432 | |
CFCC 52765 | China | Castanea mollissima | Fagaceae | MH542484 | MH542640 | MH545391 | MH545433 | |
CFCC 52766 | China | Quercus aliena var. acutiserrata | Fagaceae | MH542485 | MH542641 | MH545392 | MH545434 | |
Dendrostoma qinlingense | CFCC 52732* | China | Quercus wutaishanica | Fagaceae | MH542471 | MH542633 | MH545378 | MH545420 |
CFCC 52733 | China | Quercus aliena var. acutiserrata | Fagaceae | MH542472 | MH542634 | MH545379 | MH545421 | |
Dendrostoma quercinum | CFCC 52103* | China | Quercus acutissima | Fagaceae | MG682077 | MG682017 | MG682037 | MG682057 |
CFCC 52104 | China | Quercus acutissima | Fagaceae | MG682078 | MG682018 | MG682038 | MG682058 | |
CFCC 52105 | China | Quercus acutissima | Fagaceae | MG682079 | MG682019 | MG682039 | MG682059 | |
Dendrostoma quercus | CFCC 52734 | China | Quercus sp. | Fagaceae | MH542473 | – | MH545380 | MH545422 |
CFCC 52735 | China | Quercus sp. | Fagaceae | MH542474 | – | MH545381 | MH545423 | |
CFCC 52736 | China | Quercus sp. | Fagaceae | MH542478 | – | MH545385 | MH545427 | |
CFCC 52737 | China | Quercus sp. | Fagaceae | MH542475 | – | MH545382 | MH545424 | |
CFCC 52738 | China | Quercus sp. | Fagaceae | MH542477 | – | MH545384 | MH545426 | |
CFCC 52739* | China | Quercus sp. | Fagaceae | MH542476 | MH542635 | MH545383 | MH545425 | |
CFCC 52740 | China | Quercus sp. | Fagaceae | MH542479 | – | MH545386 | MH545428 | |
Dendrostoma shaanxiense | CFCC 52741* | China | Castanea mollissima | Fagaceae | MH542486 | MH542642 | MH545393 | MH545435 |
CFCC 52742 | China | Castanea mollissima | Fagaceae | MH542487 | MH542643 | MH545394 | MH545436 | |
Dendrostoma shandongense | CFCC 52759* | China | Castanea mollissima | Fagaceae | MH542504 | MH542652 | MH545411 | MH545453 |
CFCC 52760 | China | Castanea mollissima | Fagaceae | MH542505 | MH542653 | MH545412 | MH545454 | |
Disculoides eucalypti | CBS 132183* | Australia | Eucalyptus sp. | Myrtaceae | JQ685517 | JQ685523 | MH545413 | MH545455 |
Disculoides eucalyptorum | CBS 132184* | Australia | Eucalyptus viminalis | Myrtaceae | JQ685518 | JQ685524 | MH545414 | MH545456 |
Microscopic observations were made in tap water except where noted. Morphological analyses of microscopic characters were carried out as described by
Ascospore isolates were prepared and grown on 2% corn meal dextrose agar (CMD; CMA: Sigma, St Louis, Missouri; supplemented with 2% (w/v) D(+)-glucosemonohydrate) or 2% malt extract agar (MEA; 2% w/v malt extract, 2% w/v agar-agar; Merck, Darmstadt, Germany). Cultures are illustrated in Figure
The newly generated sequences were aligned with the sequences of
Maximum parsimony (MP) analyses were performed with PAUP v. 4.0a165 (
Maximum likelihood (ML) analyses were performed with RAxML (
Of the 4194 characters included in the phylogenetic analyses, 703 were parsimony informative (133 from the SSU-ITS-LSU, 247 from rpb2, 323 from tef1). MP analyses revealed eight MP trees 1552 steps long, one of which is shown as Figure
Phylogram showing one of 8 MP trees 1552 steps long revealed by PAUP from an analysis of the combined ITS-LSU-rpb2-tef1 matrix of Dendrostoma, with Chrysocrypta corymbiae, Disculoides eucalypti and D. eucalyptorum added as outgroup taxa. MP and ML bootstrap support above 50% are given above or below the branches. The asterisk (*) denotes the node collapsed in the strict consensus of the eight MP trees. Accessions in bold were sequenced in the present study; accessions in blue were isolated from Castanea, those in green from Quercus, in orange from Malus and in red from Osmanthus.
Dendrostoma mali X.L. Fan & C.M. Tian.
Sexual morph: pseudostromata immersed in bark and erumpent, causing a pustulate bark surface, consisting of an ectostromatic disc and entostroma with embedded ascomata. Ectostromatic disc flat or convex, surrounded by bark flaps. Entostroma light-coloured, prosenchymatous to nearly pseudoparenchymatous, mixed with bark cells, sometimes forming a more-or-less conical central column beneath the disc. Stromatic zones lacking or sometimes bark dorsally darkened. Ascomata perithecial, subglobose. Ostioles flat in the disc or slightly projecting, cylindrical, often with conical apical part. Paraphyses deliquescent. Asci oblong, fusoid, narrowly clavate or subellipsoid, with a refractive apical ring, containing (4–)6–8 ascospores in various arrangements, becoming detached at maturity. Ascospores hyaline, ellipsoid, fusoid, oblong to subacicular, often inequilateral, straight to curved, bicellular, more-or-less constricted at the median or eccentric septum, smooth, with 2–4 drops or multiguttulate, often with gelatinous terminal appendages. Asexual morph: conidiomata acervular, either forming lateral locules on the ostiolar level of sexual pseudostromata or separate, conical to pulvinate, immersed-erumpent from bark; wall pseudoparenchymatous. Often a pseudoparenchymatous conical central column present beneath the covering layer. Conidiophores non-differentiated, hypha-like or reduced to conidiogenous cells. Conidiogenous cells phialidic, lining the inner walls of cavities, subcylindrical to ampulliform, hyaline, shades of brown with age. Conidia hyaline, aseptate, smooth, multiguttulate or not, thin-walled, oblong, ellipsoid to fusoid, straight or curved.
Dendrostoma atlanticum is recognized by clay-coloured ectostromatic discs and ascospores having large guttules and bristle-like appendages.
France, Bretagne, Dépt. Morbihan (56), Saint Martin sur Oust, Beauvais, on twigs of Castanea sativa, soc. immature Valsaria sp., 15 Jan. 2016, A. Delannoy (
Atlanticum, referring to its occurrence in the Atlantic region.
Sexual morph: pseudostromata 1–4.5 mm in their widest dimension in cross section, bluntly conical or pulvinate, circular, elliptic or irregular in outline, scattered, gregarious to confluent up to 7 mm length. Ectostromatic discs 0.4–2 mm in their widest dimension, distinct and conspicuous, projecting up to 0.5(–1) mm from the bark surface, pulvinate, circular, angular or fusoid in outline, with flat or convex top, initially whitish, turning pale to dark clay-coloured, splitting the periderm, often surrounded by bark flaps. Ostioles 1–40 per disc, often originating eccentrically from the perithecial venter, arranged in ring-like configuration or variably filling the disc, (44–)100–163(–195) µm (n = 42) in diameter at the tip, brown to black, cylindrical, sometimes attenuated towards tip, plane with the disc or projecting up to 300 µm; tip usually with dark umbilicate centre. Entostroma whitish, yellowish to pale bark coloured, consisting of thin-walled, hyaline to subhyaline 1–3 µm wide hyphae and bark cells. Perithecia (390–)480–660(–750) µm (n = 35) in diameter, depressed subglobose, collapsing upward upon drying; peridium ca 10–30 µm thick, colourless to pale olivaceous, consisting of hyaline to yellowish or pale brownish, thick-walled cells without clear contours, smaller and more-or-less isodiametric outside, larger and compressed inside, very variable, (3–)4–17(–38) in diameter (n = 66). Paraphyses of broad collapsing threads. Asci (64–)71–86(–90) × (11–)13–17(–19) µm (n = 35), fusoid to oblong, being released at maturity, containing 8 biseriate ascospores. Ascospores (13–)15–18(–20) × (4.3–)5.5–7(–8) µm, l/w (2.1–)2.4–2.9(–3.9) (n = 51), ellipsoid, often inequilateral, 2-celled, slightly constricted at the median septum, with the upper cell often slightly wider than the lower, hyaline, with 1–2 large and several small guttules per cell, smooth, with a hyaline, bristle-like, straight to curved appendage (10–)11.5–15.5(–21) × (1.5–)2–2.5(–2.8) µm (n = 101) at each end.
Asexual morph acervular. Conidiomata ca 1–2.2 mm in diameter, bluntly conical, width exceeding height, prosenchymatous. Covering discs 0.3–1.1 mm in diameter, flat to pulvinate, whitish cream to pale reddish brown. Central column whitish to reddish brown, usually darker toward the top; fertile chamber ring-like around the central column; walls and column consisting of pale yellowish brown textura angularis, outer wall and outer layer of the column containing numerous crystals. Phialides (3.7–)6.3–9.7(–11.5) × (2–)2.5–3.8(–4.7) µm (n = 46), arranged in a palisade on hyaline to yellowish, angular cells, ampulliform to lageniform, less commonly cylindrical. Conidia 1-celled, hyaline, smooth, dimorphic, both morphs formed in the same locule, either ellipsoid to oblong, (6.4–)7.7–10.2(–11.7) × (4–)4.5–5.7(–6) µm, l/w (1.4–)1.4–2.2(–3) (n = 21), with a large guttule and often distinct abscission scar, or cylindrical, (7.7–)10.2–13.5(–15.3) × (2.3–)2.5–3.2(–3.5) µm, l/w (2.8–)3.6–4.7(–5.6) (n = 45), straight or curved, with mostly 3 or 4 confluent guttules.
On CMD at 16 °C in the dark colony more-or-less circular, of loose mycelium, first white, variably covered by white aerial hyphae, becoming dense, forming white and apricot to orange zones, darkening and turning black from the centre, sometimes forming reddish brown dots, spots or tubercles.
Spain, Galicia, Pontevedra, O Grove, 42°28'04"N, 08°53'14"W, on twigs of Castanea sativa, 4 Nov. 2018, M.A. Delgado (
Dendrostoma atlanticum is easily recognized by its long-pedicellate ascospores having 2–4 large drops, setting it apart from D. castaneum, which has narrow, often curved ascospores with small drops and short appendages. All species described from Castanea in China are only known from asexual morphs (
Dendrostoma atlanticum. Sexual morph a–d ectostromatic discs and ostioles e pseudostroma in vertical section f pseudostroma in cross section g peridium in cross section (in 3% KOH) h–k asci l–r ascospores. a–c, f, h, i, k–p
Dendrostoma atlanticum (
Valsa castanea Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 202 (1863) (Basionym).
≡ Amphiporthe castanea (Tul. & C. Tul.) M.E. Barr, Mycol. Mem. 7: 142 (1978).
≡ Cryptodiaporthe castanea (Tul. & C. Tul.) Wehm., Trans. Br. mycol. Soc. 18(4): 284 (1934) [1933].
≡ Diaporthe castanea (Tul. & C. Tul.) Sacc., Syll. fung. (Abellini) 1: 624 (1882).
= Cryptospora leiphaemoides Fuckel, Jb. nassau. Ver. Naturk. 25–26: 323 (1871).
≡ Diaporthe leiphaemoides (Fuckel) Sacc., Syll. fung. (Abellini) 1: 624 (1882).
Dendrostoma castaneum is recognized by KOH+ purple ostioles, slender ascospores with small drops and subfusiform conidia, and the presence of hyphal conidiophores.
Sexual morph: pseudostromata 0.8–3(–5) mm in their widest dimension in cross section, very variable, flat subconical or lenticular, in outline circular, elliptic or elongate, scattered, gregarious or confluent, and forming elongate patches, lifting the periderm slightly and often becoming visible as a dark zone on the bark surface, causing bumps in bark, splitting the periderm. Ectostromatic discs 0.3–2.7 mm in their widest dimension, often ill-defined and variable, cream, yellowish brown to dark brown, flat, surrounded by bark flaps, first present as a covering layer with ostiolar necks subsequently bursting through it, soon crumbling away. Ostioles 1–25 per disc, usually arising eccentrically from the perithecial venter, (53–)71–125(–180) µm (n = 51) in diameter, bluntly conical or cylindrical with black sides and red, yellowish, or greenish tip, often attenuated to a minute, ca 20–40 µm wide dark centre, in section rounded to angular, sometimes sulcate, variably arranged in the disc, projecting to 0.2 mm, periphysate; red colour of the ostiolar tip turning purple in 3% KOH and yellow in lactic acid. Entostroma yellowish to shades of brown, consisting of bark cells and hyaline to yellowish, 1.5–4.5 wide, thin-walled hyphae becoming thicker-walled and forming a pseudoparenchyma in the vicinity of perithecia. Perithecia tightly aggregated, (265–)305–460(–600) µm (n = 47) in diameter, depressed subglobose to ellipsoid, collapsing upward; peridium ca 10–30 µm thick, hyaline, pale olivaceous to brown, in section outside of brown isodiametric to strongly compressed thick-walled cells, inside of compressed and elongated hyaline to brownish cells, in combination (3–)4–15(–28) µm (n = 57) in diameter. Paraphyses absent at maturity. Asci (49–)53–63(–65) × (7.8–)8.5–10.5(–12) µm (n = 35), narrowly clavate to subfusoid or oblong, floating freely in the centre, thick-walled at the apex containing a minute refractive ring invisible in 3% KOH, containing 4–8 biseriate ascospores. Ascospores (11.5–)14–18(–20) × (3–)3.5–4.5(–5.3) µm, l/w (2.7–)3.5–4.6(–5.4) (n = 76), 2-celled, not or slightly constricted at the median or slightly eccentric septum, oblong to inequilaterally ellipsoid, straight to mostly curved, with the upper cell often slightly wider than the lower, broadly rounded at the ends, hyaline, with several minute drops (confluent to 2 larger drops per cell in mounts), smooth, with or without a hyaline, subconical to filiform appendage (2.2–)2.8–4.5(–5.5) × (1.1–)1.3–1.6(–1.8) µm (n = 88) at each end invisible in 3% KOH.
Asexual morph co-occurring with the sexual morph, acervular, pulvinate, scattered to aggregated, 0.5–2.7 mm in diameter, appearing as superficial discs 0.3–2 mm in diameter, with undulate surface, cream to pale brown and becoming brittle in the centre and nearly black at the periphery and often also indicated as dark zone on the bark surface around the disc; inside consisting of a pale or yellowish brown, loose and brittle central column consisting of pale brown t. prismatica and a lateral ring-like, dense, white to distinctly yellow fertile part with even or undulating margin, the latter also raising above the column, outside surrounded by a partly undulating, ca 20–25 µm thick black wall consisting of dark brown textura angularis of cells 4–10 µm in diameter at apical and upper peripheral regions, becoming paler downward and being absent at the base and lower sides. Interior of the fertile chambers consisting of isodiametric to elongate hyaline supporting cells and richly and irregularly branched hyphal conidiophores bearing phialides and conidia. Wall, supporting cells and phialides turning dilute violaceous in 3% KOH. Phialides arranged on supporting cells in palisades along the walls and on conidiophores, (6–)8.2–12(–15.3) × (1.7–)2.5–3.5(–5) µm (n = 80), repetitive, mostly lageniform, often with long necks; conidia also formed on cylindrical pegs and denticles. Conidia (6–)6.7–8(–8.8) × (2.5–)3–3.5(–3.7) µm, l/w (1.7–)2.1–2.6(–3.1) (n = 85), subfusiform, subclavate or ellipsoid, scar often distinct, smooth, with few minute drops.
On CMD at 16 °C in the dark colony circular, dense, white, covered by white cottony aerial hyphae, partly turning pale apricot, reverse orange, not zonate.
(all on recently detached twigs of Castanea sativa on ground). AUSTRIA, Burgenland, Forchtenstein, Kohlstatt, 13 Feb. 2016, H. Voglmayr (
Sizes of pseudostromata and acervuli strongly depend on twig thickness. Remarkably, red colour of the ostiolar tip, when present, turns purple in 3% KOH and yellow in lactic acid, a feature, which is typical of the Hypocreales and within the Diaporthales otherwise only found in the Cryphonectriaceae.
So far, confirmed records of D. castaneum are only known from Europe where the species is widely co-occurring with its host, Castanea sativa.
Dendrostoma castaneum. Sexual morph a–d ectostromatic discs and ostioles (in a ostioles breaking through covering layer) e pseudostroma in vertical section f pseudostroma in cross section g peridium in section (in 3% KOH) h–k asci l–s ascospores a, c–g, j, k, s
Dendrostoma castaneum (
Dendrostoma creticum is recognized by long, subacicular ascospores.
Greece, Crete, near Askifou, 35°17'47"N, 24°12'33"E, on twigs of Quercus coccifera, soc. Cytospora (Valsa morph) sp., 6 June 2015, H. Voglmayr & W. Jaklitsch (
Creticum, referring to its occurrence, Crete.
Sexual morph: pseudostromata 0.6–1.6 mm in their widest dimension in cross section, pulvinate, circular, elliptic or irregular in outline, scattered, gregarious to confluent up to 4 mm length, causing small bumps in the bark, splitting the periderm. Ectostromatic discs 0.25–1.4 mm in their widest dimension, medium to dark brown, flat or convex, surrounded by bark flaps. Ostioles 1–7 per disc, (31–)55–102(–135) µm (n = 40) in diameter at the rounded tip, dark brown to black, bluntly conical, plane with the disc or slightly prominent. Entostroma pale bark coloured, mottled. Perithecia (245–)320–445(–495) µm (n = 30) in diameter, depressed-subglobose, collapsing upward; peridium ca 10–50 µm thick, a dark brown textura angularis in face view, in section outside of dark brown textura angularis to strongly compressed cells (4–)7–14(–18) µm (n = 30) in diameter, inside of strongly compressed and elongated hyaline cells. Paraphyses absent at maturity. Asci (66–)71–85(–94) × (8.8–)9.5–11.2(–12.3) µm (n = 44), narrowly clavate to subfusoid, floating freely in the centre, containing 8 bi- to triseriate ascospores. Ascospores (26–)33–45.5(–52) × (2.7–)3–3.7(–4.6) µm, l/w (6.8–)9.8–14.3(–17.5) (n = 40), 2-celled, slightly constricted at the median or often distinctly eccentric septum, oblong, straight to curved, with the upper cell often slightly wider than the lower, hyaline, multiguttulate, smooth, with or without a hyaline subconical appendage (1.4–)1.5–2.3(–3.2) × (0.6–)0.9–1.3(–1.5) µm (n = 25) at each end.
Asexual morph unknown.
On CMD at 16 °C in the dark colony circular to irregular, dense, white, partly covered by short, white aerial hyphae, zonate, soon turning dark brown to black with pale apricot spots and margin and apricot to orange pigment diffusing into agar, reverse dark brown with orange margin.
Dendrostoma creticum is similar to the closely related D. istriacum but differs by distinctly longer ascospores, darker ectostromatic discs and a different host species.
Dendrostoma istriacum is recognized by narrow, oblong ascospores with small drops.
CROATIA, Istria, Rovinj, near Kamp Amarin, 45°06'33"N, 13°37'02"E, on twigs of Quercus ilex, soc. Diplodia sp., 14 May 2015, H. Voglmayr (
Istriacum, referring to its occurrence, Istria.
Sexual morph: pseudostromata 0.6–1.5 mm in their widest dimension in cross section, pulvinate, circular or elliptic in outline, scattered or tightly aggregated in large numbers, causing bumps in the bark and bark lesions to ca 3.2 mm long parallel to the twig axis. Ectostromatic discs 0.15–0.7 mm in diameter, mostly inconspicuous, surrounded by bark flaps, flat or convex, prosenchymatous, first whitish, turning pale to dark brown, becoming disintegrated and replaced by black ostioles and perithecial tops. Entostroma whitish to pale bark coloured. Stromatic tissues consisting of bark cells and 2–4 µm wide, hyaline to brown hyphae. Ostioles 1–5 per disc, (45–)61–91(–103) µm (n = 30) in diameter, short cylindrical, slightly projecting from the disc, brown to black; wall consisting of dark brown textura angularis. Perithecia (230–)280–393(–443) µm (n = 20) in diameter, globose to subglobose; peridium ca 15–35 µm thick, pale olivaceous to dark brown, consisting of 2–4 cell layers of thick-walled, dark brown angular cells (3–)4–13.5(–20.5) µm (n = 40) in diameter outside and long compressed, thin-walled, hyaline to brownish cells inside. Paraphyses absent at maturity. Asci (59–)62–70(–74) × (7–)8.5–10(–11) µm (n = 30), fusoid to narrowly clavate, floating freely in the centre, containing 8 bi- to triseriate ascospores. Ascospores (19.3–)20.5–25.5(–29.5) × (3–)3.5–4.2(–5.1) µm, l/w (4.5–)5.3–7(–8.7) (n = 40), 2-celled, constricted at the more-or-less median septum, oblong, straight to curved, with the upper cell often slightly wider than the lower, hyaline, containing several small guttules concentrated towards the ends and the septum, smooth, with a hyaline subconical appendage (1.7–)2.5–3.5(–4.5) × (0.8–)1–1.3(–1.5) µm (n = 40) at each end, becoming elongated in mounts.
Asexual morph: conidiomata ca 250–520 µm in diameter, acervular, inconspicuous, immersed in bark, causing small bark bumps, becoming visible in fissures, whitish to brownish, flat or convex, bluntly conical, usually broader than high, consisting of a broad sterile greyish brown central column, a white outer fertile ring and a brown covering layer; also fertile between the latter and the top of the column. Covering layer consisting of a dark brown textura angularis of 4–10 µm wide cells, turning paler to hyaline and more rounded downwards; column comprising pale brown textura angularis-epidermoidea of similarly sized cells; outer margin of the fertile ring consisting of a narrow layer of hyaline to pale brown, angular to compressed cells; gel surrounding rounded to angular, subhyaline to hyaline cells supporting phialides slowly turning pinkish in 3% KOH. Phialides forming palisades in fertile areas, tightly packed, cylindrical to ampulliform, often with long acute necks, (5.5–)6.3–9(–11) × (1.8–)2.2–3.7(–5.3) µm (n = 33). Conidia (4–)5–6.6(–7.4) × (1.9–)2.1–2.5(–2.7) µm, l/w (1.6–)2.1–3(–3.7) (n = 53), oblong to ellipsoid, 1-celled, hyaline, smooth, usually with distinct abscission scar.
On CMD at 16 °C in the dark, colony circular to irregular, dense, white, partly covered by short white aerial hyphae, zonate, soon turning dark brown to black with pale apricot to reddish brown spots and margin and some pale apricot pigment diffusing into agar, reverse dark brown with pale apricot margin.
CROATIA, Istria, Rovinj, near Kamp Veštar, 45°03'19"N, 13°40'55"E, on twigs of Quercus ilex, 30 May 2019, H. Voglmayr (
Dendrostoma istriacum is closely related to D. creticum but differs from that species by distinctly shorter ascospores and a different host species.
Dendrostoma istriacum (
Sphaeria leiphaemia Fr., Syst. mycol. (Lundae) 2(2): 399 (1823) (Basionym).
≡ Amphiporthe leiphaemia (Fr.) Butin, Sydowia 33: 22 (1980).
≡ Diaporthe leiphaemia (Fr. : Fr.) Sacc. [as ‘leiphaema’], Atti Soc. Veneto-Trent. Sci. Nat. 2(1): 135 (1873).
≡ Valsa leiphaemia (Fr.) Fr., Summa veg. Scand., Sectio Post. (Stockholm): 412 (1849).
Dendrostoma leiphaemia is recognized by conspicuous ectostromatic discs, broad conical ostiolar necks, and broad multiguttulate ascospores.
Sexual morph: pseudostromata 1–5 mm in their widest dimension in cross section, pulvinate to conical, circular, elliptic or irregular in outline, scattered, aggregated to confluent, sometimes forming lines of up to 15 mm length, causing conspicuous bumps and lesions in the bark; dark brown dorsal zones present within the bark, absent in basal regions. Ectostromatic discs 0.35–2.5 mm in their widest dimension, conspicuous, whitish, cream, pale brown, pale yellowish brown to dull brown, fusoid, triangular to circular in section, flat or convex, often surrounded by bark flaps, elevated up to 1.3 mm beyond the bark surface, brittle to powdery, first present as a covering layer with ostiolar necks subsequently bursting through it, eventually crumbling away. Ostioles 1–30 per disc, (88–)124–220(–336) µm (n = 64) in diameter, dark brown, black, or reddish brown with black, rarely yellowish tip, cylindrical with conical apical part, attenuated to 35–90(–180) µm at the rounded, compressed or coarsely sulcate tip, projecting to 250, less commonly 400 µm, white, in upper regions sometimes yellow inside, periphysate, arising centrally to eccentrically from the perithecial venter and slightly convergent above perithecia; turning partly yellow, partly brown in 3% KOH. Entostroma whitish to pale yellowish or pale bark-coloured, prosenchymatous to pseudoparenchymatous, the latter particularly in the vicinity of perithecia, consisting of 1.5–5 µm wide hyphae or angular cells, mixed with bark cells. Perithecia arranged in valsoid configuration, tightly aggregated, (292–)380–625(–700) µm (n = 21) in diameter, globose to depressed-subglobose, with gelatinous contents, collapsing upward; peridium ca 7–35 µm thick, pale olivaceous to dark brown, consisting of an outer layer of isodiametric to elongate, thick-walled dark brown cells and an inner layer of compressed elongate, hyaline to brownish, thin-walled cells (5–)6.5–16(–22.5) µm (n = 31). Paraphyses absent at maturity. Asci floating freely in the centre when mature, (49–)58–71(–80) × (9–)10–13.5(–17.5) µm (n = 56), clavate, oblong, fusoid to subellipsoid, with a refractive apical ring, containing 8 bi- to triseriate, fasciculate or obliquely uniseriate ascospores. Ascospores (15–)16–19(–21) × (3.8–)4.3–5.2(–5.8) µm, l/w (2.7–)3.3–4.1(–4.7) (n = 95), 2-celled, not or slightly constricted at the median or slightly eccentric septum, inequilaterally ellipsoid or oblong, straight or curved, with the upper cell sometimes slightly wider than the lower, hyaline, multiguttulate, smooth, lacking appendages.
Asexual morph co-occurring with the sexual morph, acervular, either present as locules in lateral regions of pseudostromata above perithecia or forming separate, conical to pulvinate, dorsally blackened acervuli 0.9–2.2 mm in diameter, with conical upper part or whitish to cream or brownish, more-or-less circular, continous or deeply fissured discs ca 0.4–1 mm in diameter and whitish-cream, partly hollow interior containing slightly darker fertile chambers meandering through it. Walls and interior consisting of brown or hyaline to pale yellowish brown textura angularis. Phialides lining inner wall of the cavity, sessile, (4.8–)6.5–11(–12.7) × (1.7–)2–3.8(–5.3) µm (n = 16), subcylindrical to lageniform, reddish brown in 3% KOH (when old). Conidia (4.8–)7–9.5(–11) × (1.5–)1.8–2.3(–2.5) µm, l/w (2.3–)3.3–4.9(–6.3) (n = 50), unicellular, cylindrical, oblong, subclavate, rhomboid or narrowly ellipsoid, straight to slightly curved, often with a truncate or acute end, hyaline, turning pinkish-yellowish in 3% KOH, smooth, with minute terminal drops, adhering together in masses when old.
On CMD at 16 °C in the dark, colony irregular or dimorphic, dense, white, partly covered by short white aerial hyphae, zonate, soon turning dark brown to black with red or reddish brown spots, reverse dark brown, reddish brown with white, pale apricot or reddish brown spots and margins.
AUSTRIA, Kärnten, St. Margareten im Rosental, shrubs in front of the Stariwald, grid square 9452/4, on branches of Quercus petraea, 9 Jan. 1995, W. Jaklitsch W.J. 443 (
Asexual fructifications of this species are reported to have dimorphic conidia (
Dendrostoma leiphaemia. a–s Sexual morph a–e ectostromatic discs and ostioles f pseudostroma in cross section g pseudostroma in vertical section h peridium in cross section i, n–s ascospores j–m asci t–z asexual morph t conidioma in cross section u, v phialides w–z conidia; a, d–g, t–z
1 | Ascospores without appendages, multiguttulate, 15–21 × 4–6 µm; on broad-leaved Quercus spp. | D. leiphaemia |
– | Ascospores with appendages | 2 |
2 | Appendages 10–21 µm long, bristle-like; ascospores with 1–2 large guttules per cell, 13–19.5 × 4.5–8 µm; on Castanea sativa | D. atlanticum |
– | Appendages <6 long, not bristle-like | 3 |
3 | Ascospores 11.5–20 × 3–5.3 µm, oblong, often curved, with 2 minute guttules per cell; on Castanea sativa | D. castaneum |
– | Ascospores longer | 4 |
4 | Ascospores oblong, 19–30 × 3–5 µm; on Quercus ilex | D. istriacum |
– | Ascospores oblong to subacicular, 26–52 × 2.7–4.5 µm; on Quercus coccifera | D. creticum |
Our phylogenetic analyses are largely congruent with those of
Previous authors recorded phytopathogenic potential in all species of Dendrostoma studied by them (
Although other genera of the Erythrogloeaceae produce acervuli, asexual morphs of Dendrostoma have been termed pycnidia (
Most species of Dendrostoma are only known as asexual morphs. Only one of the 10 species described by
The high species biodiversity of Dendrostoma recorded from Eastern Asia as well as the phylogenetic patterns indicate that the group may have originated in this area. This is also supported by the fact that the European species do not form a monophyletic group, but are embedded within Eastern Asian lineages, indicating that Europe has been colonised from Asia several times independently. In addition, evolutionary radiation may have started on Castanea as the basal subclade A exclusively contains accessions from that host (Fig.
We thank Alain Delannoy, Miguel Angel Delgado, and Joseba Castillo for providing fresh material of Dendrostoma atlanticum and Irmgard Greilhuber and Walter Till (