Molecular results
The obtained sequences of the three Schroeteria decaisneana samples fully concur in the ITS region between the two apothecial isolates (teleomorph) and the sorus isolate (anamorph), except for 1 nt in the ITS1 in one of the apothecial isolates (Tab. 2). Also in the LSU D1–D2 no difference was observed between the two apothecial isolates and the sorus isolate (Tab. 2). The two S. poeltii sequences likewise fully concur in the ITS and LSU D1–D2 region between apothecial and sorus isolate, except for 1 nt in the LSU. Considering the interspecific distances among the Schroeteria species and the identity of the host species, our result proves that teleomorph and anamorph belong together.
The interspecific distances in the ITS (457–462 nt in core clade of Schroeteria) and LSU D1–D2 (598 nt) range at 5–6.3% / 2.5–2.7% within the core clade, but at ~14% / 8.5–9.6% between core clade and Schroeteria poeltii, 16.7–18% / 8.9–9.7% between core clade and Ciboria ploettneriana, and 9.2% / 3.7–3.9% between Schroeteria poeltii and Ciboria ploettneriana, respectively. The S1506 intron is absent in all sequences in which the 3'-end region of SSU was included (the region is missing in Schroeteria bornmuelleri).
Table 2. Inter- and intraspecific distance matrix of the ITS1-5.8S-ITS2 region (before slash) and LDU D1–D2 domain (after slash) for Schroeteria spp. in comparison with Ciboria ploettneriana (p-distances, transitions + tansversions, pairwise deletion).
|
S. decaisneana
|
S. delastrina
|
S. bornmuelleri
|
S. poeltii
|
Cib. ploettneriana
|
S. decaisneana
|
0–0.2 / 0 %
|
|
|
|
|
S. delastrina
|
5–5.3 / 2.5–2.7 %
|
0–0.4 / 0.2 %
|
|
|
|
S. bornmuelleri
|
6–6.3 / – %
|
5.2 / – %
|
– / –
|
|
|
S. poeltii
|
14–14.3 / 9.4–9.6 %
|
14.2–14.7 / 8.5–8.9%
|
13.8 / – %
|
0 / 0.2 %
|
|
Cib. ploettneriana
|
17.2–18 / 9.7 %
|
17.2–17.6 / 8.9–9 %
|
16.7 / – %
|
9.2 / 3.7–3.9 %
|
– / –
|
A strongly supported Schroeteria core clade is formed in our combined maximum likelihood analysis of ITS+LSU (Pl. 1), but also in separate analyses of ITS and LSU (S1, S2). The clade comprised S. decaisneana, S. delastrina, and S. bornmuelleri, and nested in our combined analysis in an unsupported clade with various Monilinia spp. of section Disjunctoriae growing on fruits of Ericaceae and Rosaceae (Pl. 1). When separately analysing the ITS region (S1), this clade was strongly supported but included also Ciborinia foliicola, C. whetzelii, and Ciboria betulae, whereas when separately analysing LSU (S2) the Schroeteria core clade clustered with medium support with Ciboria shiraiana (on Morus), which in the other two analyses formed a supported clade with C. carunculoides (also on Morus) outside Sclerotiniaceae and Rutstroemiaceae.
The Schroeteria core clade clustered particularly with two Ericaceae inhabiting species: it formed a medium (Pl. 1, S1) or strongly (S3) supported sister clade of M. jezoensis, though with a rather long branch, and together with this species a less supported sister clade of M. azaleae (Pl. 1, S1). Both species grow on Rhododendron and are members of the Monilinia alpina group of section Disjunctoriae, to which also M. cassiopes belongs (Batra 1991: 102), but apparently also Stromatinia pyrolae.
In contrast to the phylogenetic affiliation of the Schroeteria core clade with Monilinia section Disjunctoriae, Schroeteria poeltii clustered distant from Schroeteria s.str. It formed an unsupported (Pl. 1) or strongly supported (S3) distant sister clade to the supported clade of Sclerotinia bulborum and S. 'binucleata' (an undescribed species of Sclerotinia s.l. parasitizing on Ranunculus ficaria and Corydalis cava). Ciboria ploettneriana clustered with comparatively short branches with these and other typical members of Sclerotinia, Stromatinia, and Grovesinia in an unsupported clade (Pl. 1, S1).
Morphological results
In the following, Ciboria ploettneriana, Schroeteria decaisneana, and S. poeltii are described in detail from their teleomorph, the latter two also from their anamorph.
Key to species of Sclerotiniaceae with known teleomorph growing on Veronica seeds
- Ascospores mainly *12.5–16.5 × 6–7.5 µm, with several LBs 0.5–0.8 µm diam. grouped near each end, (2–)4-nucleate; living paraphyses containing ± refractive vacuolar bodies (VBs) in terminal cell; on blackened seeds of Veronica hederifolia agg., climate temperate humid ....................................................... Ciboria ploettneriana
- Ascospores mainly *10–13 × 5.5–6.5 µm, with 1(–2) LBs 1.4–2.7 µm diam. near each end, uninucleate; living paraphyses without refractive vacuoles in terminal cell .........................................................................................2
- Apothecial stipes 3–13 mm long; ascospores ellipsoid to fusoid, LBs subpolar, 1.4–2.7 µm diam.; on light to bright (grey-)brown seeds of Veronica hederifolia agg., climate temperate humid ........ Schroeteria decaisneana
- Apothecial stipes 5–25 mm long; ascospores ellipsoid, LBs more polar, 1.5–1.8 µm diam.; on blackened seeds of Veronica cymbalaria, climate mediterranean semihumid ...................................................... Schroeteria poeltii
Table 3. Comparison of teleomorph characteristics of Schroeteria decaisneana, S. poeltii, and Ciboria ploettneriana. # = data evaluated from Nagler et al. (1989).
|
Schroeteria decaisneana
|
Schroeteria poeltii
|
Ciboria ploettneriana
|
Ascospores size [µm]
|
*(9.5–)11–13(–14.5) × (5–)5.5–6.5(–6.7)
|
*(9.5–)10–11(–11.3) × (5.3–)5.5–6(–6.3)
|
*(10.5–)12.5–16.5(–19) × (5.5–)6–7.5(–9)
|
- shape
|
ellipsoid to fusoid or slightly ovoid
|
ellipsoid to slightly ovoid
|
ellipsoid- to ovoid-fusoid
|
- LBs (living state!)
|
1(–2) LBs of 1.4–2.7 µm diam.
and some small ones near each end
|
1(–2) LBs of (1–)1.5–1.8 µm diam.
and some small ones at each end
|
1(–2) LBs of 0.5–0.8(–1) µm diam.
and some small ones near each end
|
- nuclei in ascospores
|
1 nucleus, 2.8–3 µm diam.
|
1 nucleus, 3–3.2 µm diam.
|
(2–)4-nuclei, 2.2–2.9 µm diam.
|
Phialides [µm]
|
†2.5–5 × 1.5–3
|
8–20 × 3.5–5.8#
|
*6–10 × 2.6–3.5(–4)
|
- phialoconidia [µm]
|
*2.8–3.6 × 2.6–3.4
|
(2.7–)3–3.6(–4) × (2.7–)2.9–3.5(–3.8)#
|
*2.5–3.2 × 2.5–3
|
Asci [µm]
|
*(145–)157–185(–192) × 11–12.3(–13)
|
*145–160 × 9.5–10.5(–10.8)
|
*180–210 × (10–)11–12(–13)
|
Paraphysis terminal cells
|
with non-refractive vacuoles
|
with non-refractive vacuoles
|
with slightly to strongly refractive VBs
|
Ectal excipulum
|
vertically or often horizontally oriented
textura globulosa(-prismatica)
|
vertically oriented
textura globulosa(-prismatica)
|
vertically oriented
textura globulosa-prismatica
|
- lower flank cells [µm]
|
*(16–)20–45(–70) × (12–)15–28(–35)
|
*20–50 × 15–30
|
*20–40(–58) × 15–25(–30)
|
- marginal cells [µm]
|
*38–70 × 6–10
|
*25–35 × 13–15
|
*8–15(–25) × 5–11.5
|
- cells in stipe [µm]
|
*28–46 × 3–4
textura porrecta
|
†25–40 × 6–9.5
textura prismatica-porrecta
|
*20–45 × 7–17
textura prismatica
|
Apoth. disc diam. [mm]
|
1.5–3.5
|
(1–)1.3–2.5(–3)
|
(1.2–)2–4
|
- stipe colour
|
medium grey-brown, near base blackish-brown
|
ochraceous, near base blackish-brown
|
whitish to creamish, sometimes brownish below
|
- stipe size [mm]
|
(3–)5–10(–13) × (0.2–)0.4–0.6(–0.8)
|
5–18 × 0.2–0.55
|
2–7 × (0.35–)0.5–0.8(–1)
|
Seed size (infected) [mm]
|
2–2.5
|
(0.9–)1.7–2.8
|
2–3.2
|
- surface
|
light to bright (grey-)brown
|
bright grey-brown to blackish
|
blackish-brown to black
|
Phenology (apothecia)
|
XI–V
|
I
|
IV–V
|
Host
|
Veronica hederifolia agg.
|
Veronica cymbalaria
|
Veronica hederifolia agg.
|
climate
|
temperate humid
|
meso-/supramediterranean semihumid
|
temperate humid
|
Ciboria ploettneriana (Kirschst. in Rehm) N.F. Buchw., K. Vet. Landbohøjsk., Aarsskr. 32: 165 (1949) — Pls 2–6
≡ Sclerotinia ploettneriana Kirschst., in Rehm, Ascom. exsicc.: no. 1603 (1905), nom. inval., Art. 38.1(a) ICN
≡ Sclerotinia ploettneriana Kirschst., in Rehm, Annls mycol. 3(5): 411 (1905), nom. inval., Art. 38.1(a) ICN
≡ Sclerotinia ploettneriana Kirschst., Verh. Bot. Ver. Prov. Brandenburg 48: 43 (1906)
Etymology: named after the collector, the German Traugott Plöttner (1853–1923), who frequently accompanied W. Kirschstein (1863–1946) on his excursions (Kummer 2010).
Lectotype (designated here, MBT 10000619): Germany, Brandenburg, Groß Behnitz, Hasellake, on black stromatized seeds of Veronica hederifolia agg., 27.IV.1905, W. Kirschstein (B 70 0100006).
DESCRIPTION. Teleomorph: Apothecia fresh (1.2–)2–4 mm diam., non-gelatinous, disc light to bright ochraceous-brown, slightly concave, finally flat, darker brown with age, margin thin, not protruding, smooth to very finely whitish denticulate, exterior pale to light ochraceous, smooth to slightly hairy, receptacle at base 0.7 mm thick, at margin 0.35 mm; stipe 2–7 × (0.35–)0.5–0.8(–1) mm, whitish to pale cream, at base often darker brown, usually somewhat hairy and with adhering particles, emerging singly (rarely also in pairs) from concave or convex side of the seed. Asci *180–210 {2} × (10–)11–12(–13) µm {3}, †150–173 × 8–10(–11) µm {1}, protruding *5–30 µm beyond paraphyses or (†) ± equalling the paraphyses, with 8 equal-sized spores (in overmature apothecia also unequal-sized), pars sporifera *61–73 µm long, spores obliquely (sub)biseriate, †65–83 µm, spores subbiseriate to uniseriate; apex (†) hemispherical to moderately truncate, apical thickening immature or mature †2–3.8 µm thick, apical ring blue in IKI {4}, euamyloid (BB), of Sclerotinia-type, lower ring strongly blue, upper parts less so; base with short and thick stalk, arising from croziers {3}. Ascospores *(10.5–)12.5–16.5(–19) × (5.5–)6–7.5(–9) µm {4}, †(11–)12–15(–16) × (4.5–)4.8–6 µm {5}, ellipsoid-fusoid to ellipsoid-fusiform (homopolar) or often distinctly ovoid-fusoid (heteropolar), ends obtuse to subacute, ± equilateral; containing a few minute LBs near each end, (0.2–)0.5–0.8(–1)((–2)) µm diam. {4}, OCI 0.5–1(–2), with ((1–))(2–)4 nuclei in centre {4}, without distinct glycogen; surrounded by a very delicate sheath that slips off the spore after ejection, spore wall surface CRB–; overmature spores 0–1(–2)-septate (septum median or sometimes eccentrical when 1-septate), *17–19 × 7.5–9 µm, without LBs; forming germ tubes and/or phialides. Paraphyses apically uninflated to slightly clavate or lageniform, terminal cell *(20–)42–100(–133) × (3–)4–5(–5.7) µm {3}, †2.5–4 µm wide {1}, lower cells *17–39 × 2.2–3.5 µm {2}; branched in lower and middle part; terminal cell containing hyaline, slightly to strongly refractive vacuolar bodies (VBs) in (11–)15–50(–70) µm long upper part {3}, VBs staining turquoise in CRB, in H2O turning light purplish-pink with age (vital state) {2}, without SCBs; when senescent ensheathed by an ochre-brass-brown exudate along entire length. Subhymenium light ochre, 30 µm thick, of dense textura intricata. Medullary excipulum subhyaline, non-gelatinized, at base of receptacle ~200–400 µm thick, near margin 100 µm, upper part of ± loose t. intricata, individual cells *40–110 × 6–16 µm, smooth, without exudate; lower part of dense, horizontal t. porrecta, 150 µm thick at base of receptacle, sharply delimited from ectal excipulum, in stipe of t. porrecta. Ectal excipulum ± hyaline, 50–150 µm thick at lower flanks, of thin-walled (wall †0.2–0.4 µm thick), ± vertically oriented t. globulosa-prismatica, cells *20–40(–58) × 15–25(–30) µm {2}, cortex of smaller globose cells which sometimes form clavate to stalked hair-like protuberances of *10–21 × 8–10(–11) µm; at mid flanks 30–40 µm thick, at margin 20 µm thick, with ochre-brown exudate, marginal cells *8–15(–25) × 5–11.5 µm {2}, ellipsoid to clavate or stalked; exterior without hyphal layer, at lower flanks with hair-like, globose to clavate or stalked cells of *10–21 × 8–10(–11) µm, often containing VBs; in stipe of large-celled, subhyaline t. prismatica (cells *20–45 × 7–17 µm), covered by a thin layer of hyphoid, *3–7 µm wide elements, with scattered, indistinctly protruding, irregularly clavate to stalked, thin-walled, hair-like cells of *13–28 × 6–8 µm that contain VBs, at very base of stipe of dark red-brown t. angularis. Rhomboid crystals abundantly present in medullary excipulum {5}, diagonal diameter 8–22(–32) µm, small scattered crystals occurring on ectal excipulum and hymenium, crystals present also in medullary excipulum of basal part of stipe and a few in mycelial tissue within seed, sometimes forming druses 10–20 µm diam. Amyloidity of tissue: subhymenium {2} and outer medullary excipulum {2} pale to distinctly bluish in IKI (sometimes only visible after squashing). – Anamorph: Cultural characteristics: In pure culture on MEA the ascospores tardily germinated but did not form a mycelium. Smut-like synanamorph unknown. Microconidial synanamorph formed on ascospores germinating in senescent apothecia, producing phialides of *6–10 × 2.6–3.5(–4) µm that emerge terminally or laterally, either directly or on short to long germ tubes; phialoconidia globose to subglobose, *2.5–3.2 × 2.5–3 µm {3}, smooth, with a single eccentrical LB 1.2–2 µm diam.
Habitat: on fallen, heavily stromatized, previous year's seeds of Veronica hederifolia agg. {20/9}, seeds 2–3.2 mm diam., hard, surface ± rugose, entirely blackish-brown to black due to abundant, dark brown intracellular hyphae in epidermial layer and more sparse in outer parenchym cells. Desiccation tolerance: not tested for apothecia (probably intolerant), but surviving min. 9 months within the dry stromatized seeds. Phenology: IV–V. Altitude: 30–195 m. Geology: on basophilic to calcareous or more acidic soil: Muschelkalk (partly covered by loess), pleist- & holocene marl, sand & gravel, and fluviatile sediments.
Phenology of Ciboria ploettneriana
|
|
Feb-2
|
Mar-1
|
Mar-2
|
Apr-1
|
Apr-2
|
May-1
|
May-2
|
teleomorph
|
–
|
–
|
–
|
9
|
9
|
2
|
–
|
Specimens included (all on fallen seeds of Veronica hederifolia agg.).
Teleomorph: Germany: Mecklenburg-Vorpommern, 3.7 km S of Wismar, 1.2 km NE of Metelsdorf, slope at road, MTB 2134/232, 30 m, 14.IV.2019, T. Richter, vid. M. Reul (T.R., M.R. 6806). – Berlin, 8.5 km SE of Berlin, Treptow, Baumschulenweg, Späth-Arboretum, 37 m, MTB 3546/24, 30.IV.1992, D. Benkert, vid. V. Kummer (B 70 0009941, d.v.). – Brandenburg, 22.5 km NE of Brandenburg, SW of Groß Behnitz, Hasellake, 35 m, MTB 3442/123, 14.IV.1905, W. Kirschstein, vid. V. Kummer (B 70 0100005). – ibid., 27.IV.1905, W. Kirschstein, vid. V. Kummer (B 70 0100006, lectotype). – 1.7 km SW of Teupitz, ~1.5 km SE of Egsdorf, between Mittel-Mühle and Tornower See, ~45 m, MTB 3847/43, 27.IV.1972, D. Benkert, vid. V. Kummer (B 70 0009940, d.v.). – Sachsen-Anhalt, 2.2 km SE of Merseburg, WSW of Trebnitz, Werder canal, MTB 4638/31, 88 m, 20.VI.2003, G. Hensel (blackened seeds without apothecia, deposited at the same day in a box in the garden of M. Huth); 3.IV.2004, M. Huth (apothecia developed in box, M.H.). – ibid., 30.IV.2019, G. Hensel (blackened seeds without apothecia, H.B. 10210). – 4.3 km NE of Freyburg, W of Zeuchfeld, above vineyard, MTB 4736/42, 195 m, 13.IV.2009, P. & S. Rönsch (P.R., H.B. 9037). – ibid., 15. & 23.IV.2010, U. Richter. – 1.7 km ENE of Freyburg, NW-corner of Alte Göhle, Bauernholz, 185 m, MTB 4736/4, 2.V.2003, M., W. & E. Huth (W.H.). – ibid., 20.VI.2003, M. Huth & P. Rönsch (blackened seeds without apothecia, deposited at the same day in a box in the garden of M. Huth); 15. & 18.IV.2004, M. Huth & P. Rönsch (apothecia developed in box, M.H. & P.R.). – ibid., 15.IV.2007, P. & S. Rönsch (P.R., sin. doc.). – ibid., 22.IV.2010, U. Richter et al. (WU 43984, sq.: ITS/LSU MZ048354). – ibid., 23.IV.2010 U. Richter (H.B. 9271 ø). – 5.5 km WSW of Freyburg, S of Hirschroda, Hirschrodaer Graben, 185 m, MTB 4836/1, 12.IV.2004, P. Rönsch, S. Thieme, M. Huth (P.R.). – 4.3 km NW of Naumburg, 1 km WNW of Roßbach, Scherbitzberg-Schlucht, MTB 4836/2, 195 m, 8.IV.2009, W. Huth (ø). – ibid., 2.V.2019, W. Huth (W.H.).
Taxonomic remarks: Ciboria ploettneriana resembles Schroeteria decaisneana in most of its morphological traits, including various microscopic details, such as size of asci and paraphyses, size and shape of medullary and ectal excipular cells, IKI-reaction of ascus apex and medullary excipulum, and presence of crystals. The species deviates macroscopically in light-coloured, often whitish, somewhat shorter apothecial stipes with a lighter colour at the base, and seeds with a more blackish surface, causing a strong optical contrast between stipe and seed, in contrast to grey-brown seeds in S. decaisneana. Microscopically, Ciboria ploettneriana deviates from Schroeteria decaisneana in distinctly larger, slightly more heteropolar ascospores with often pointed, subacute ends. Under vital study, Ciboria ploettneriana differs in ascospores with a distinctly lower lipid content composed of much smaller LBs and (2–)4 nuclei instead of only one, and in ± strongly refractive vacuoles (VBs) in the terminal cells of paraphyses (Tab. 3). Less evident or possibly variable features of C. ploettneriana are slightly longer asci, slightly smaller conidia formed on longer phialides, and an ectal excipulum of vertically oriented cells in the receptacle (in S. decaisneana often horizontal) and wider cells in the stipe (textura prismatica vs. t. porrecta), also with much shorter cells at the margin (Tab. 3).
Variation: Ascus and spore size was rather consistent among the specimens studied. Also the size of LBs never exceeded 1 µm diam., with rare exceptions of single spores. The number of nuclei in the spores varied between two and four within all four samples in which living spores have been studied in detail (H.B. 9037 & 9271, M.R. 6806, 2.V.2019), but the tetranucleate spores were often distinctly more numerous than the binucleate spores when regarding only mature, freshly ejected spores.
Cultural studies: Microconidia were abundantly produced from the ascospores in senescent apothecia after incubation in a moist chamber. When shot on agar medium (MEA), the ascospores did not produce a mycelium even after several weeks of incubation at room temperature.
In two of the here studied samples, apothecia were obtained after incubation of blackened seeds placed in June directly . In this the apothecia developed during April of the following year.
Nomenclature and misidentifications: Hein & Gerhardt (1981) listed four voucher specimens as syntypes of Sclerotinia ploettneriana in B (Botanisches Museum Berlin-Dahlem): one from Rathenow (29.X.1899), two from Groß Behnitz (14. & 27.IV.1905), and one containing a mixture taken from both sites (29.X.1899 and 'April 1905'). Kirschstein's protologue mentions both sites with the dates 29.X.1899 and IV.1905 (without specifying the exact date in April). Under the latter date Kirschstein distributed the fungus as an exsiccata in Rehm: Ascom. exsicc. no. 1603. In the present study all four specimens, which now bear the herbarium numbers B 70 0100003–0100006, were examined by one of us (V. Kummer).
The first report of Sclerotinia ploettneriana in Rehm (1905 no. 1603, in sched.) and also the publication of this exsiccata in Annales Mycologici (Rehm 1905) is invalid, because only the collection data was provided but no description. Valid publication with description was done in Kirschstein (1906). The unillustrated protologue reports apothecia 2–3 mm diam., stipes 1–10 × 0.5 mm, emerging singly or up to four from one seed, asci 160–180 × 10–12 µm, and oval spores of 15–18 × 6–7 µm with 1–2 small oil drops. Kirschstein emphasized the black stromatization of the seeds in contrast to the whitish uninfected seeds.
The Rathenow specimen (B 70 0100003) contains principally about ten light brown seeds of Veronica hederifolia agg. from which apothecia with ellipsoid spores of †9–11 × 5–6 µm emerge (Pl. 13). The light seed colour and the small spores indicate that this collection represents Schroeteria decaisneana, although the spores often contained more than one comparatively small oil drop in each half. A single seed without adhering apothecium appears to belong to Ciboria ploettneriana because of its black-brown colour. The date of this collection (end of October) coincides with the phenology of the S. decaisneana teleomorph which comprises autumn, winter, and spring, according to our study.
The two samples from Groß Behnitz (B 70 0100005–6, one shown on Pl. 6: 1) each contain numerous bright to dark black-brown seeds of Veronica hederifolia agg., and apothecia which much better fit the protologue of Sclerotinia ploettneriana regarding spore shape (ellipsoid to fusoid, rarely clavate) and spore size (B 70 0100005: †11–16 × 5–6 µm, B 70 0100006: †12–15 × 4.5–6 µm, examined in water). Because of the small spore size in the Rathenow collection we conclude that the protologue derives from the large-spored species alone and not from a mixture with Schroeteria decaisneana. Therefore, we here designate one of the two specimens from Groß Behnitz, B 70 0100006, as lectotype of Sclerotinia ploettneriana.
The examined duplicate of Rehm's exsiccata (B 70 0100004) provided a surprize: it does not contain any seeds of Veronica, but half a dozen of sclerotia with apothecia of an undetermined Sclerotiniaceae, also a few seeds of a ?Stellaria. The original label indicates that this exsiccata was composed of a mixture of two collections: the vast majority of the material was from Groß Behnitz (IV.1905), whereas a sparse minority came from Rathenow (29.X.1899). Other duplicates of this exsiccata might therefore actually contain seeds of Veronica with apothecia of Ciboria ploettneriana.
Ecology: Ciboria ploettneriana was recorded near Freyburg at three sites, in a thermophilous deciduous forest with Ulmus minor bordered by Prunus spinosa (Zeuchfeld, Pl. 12), in a thermophilous Querco-Carpinetum (Alte Göhle), and in a more shady Aceri-Fraxinetum (Hirschrodaer Graben). The site near Merseburg was a ?Pruno-Fraxinetum river bank forest with Prunus padus (Elster-Luppe-Aue), that near Naumburg a nitrophilous Quercus-Fraxinus forest edge (Alliarion, Pl. 5), and that near Wismar a narrow forest strip of Acer, Aesculus and Tilia between road and farmland. Monilinia johnsonii on Crataegus fruits occured together with Ciboria ploettneriana at the site near Naumburg.Besides the collections from Mecklenburg-Vorpommern, Brandenburg (including Berlin), and Sachsen-Anhalt as listed under Specimens included, no further unequivocal records of Ciboria ploettneriana came to our notice.
Schroeteria decaisneana (Boud.) De Toni, in Saccardo, Syll. fung. 7(2): 501 (1888) — Pls 7–14
≡ Thecaphora decaisneana Boud., Bull. Soc. mycol. Fr. 2: 167 (1886)
≡ Geminella decaisneana (Boud.) Boud., Bull. Soc. mycol. Fr. 3(2): 150, pl. 15 fig. 2 (1887)
≡ Schizonella decaisneana (Boud.) Thirum. & M.D. Whitehead, Am. J. Bot. 55: 186 (1968)
= Schroeteria parvispora (Bref.) Ferd. & Winge, Dansk bot. Ark. 2(1): 4 (1914)
≡ Geminella parvispora Bref., Unters. Gesamtgeb. Mykol. 15: 75 (1912)
Etymology: named after the French botanist Joseph Decaisne (1807–1882), who made the first record.
Type: collected in 1868 by J. Decaisne in the surroundings of Versailles, and later during several years in spring and summer by E. Boudier near Montmorency (Paris), always on "Veronica hederacea" (= V. hederifolia agg.). No lectotype appears to have been designated, and no material was ordered in this study.
DESCRIPTION. Teleomorph: Apothecia fresh 1.5–3.5 mm diam., non-gelatinous, disc light ochraceous-brown, slightly concave, finally flat to medium convex, darker brown with age, margin thin, not protruding, ± smooth, sometimes deeper red-brown, exterior pale ochraceous, finely rough, receptacle at base 0.5–0.75 mm thick, at margin 0.25–0.35 mm; stipe (3–)5–10(–13) × (0.2–)0.4–0.6(–0.8) mm, concolorous with receptacle or darker red-brown, especially in lower part, black-brown at base, here or also upwards with ± appressed hairs; one or sometimes 2–3(–4) stipes emerging from concave or convex side of the seed. Asci *(145–)157–185(–192) × 11–12.3(–13) µm {2}, †118–147 × (7.5–)8.5–10(–11.7) µm {2}, protruding *25–35 µm beyond paraphyses or †10 µm shorter up to 10 µm longer than paraphyses, containing 8 equal-sized spores, pars sporifera *(48–)52–65 µm long, spores obliquely (sub)biseriate, †48–72 µm, spores uniseriate; apex (†) hemispherical to subtruncate or obtuse, apical thickening †1.7–2.8 µm thick when ± mature, apical ring medium to strongly blue in IKI {5}, euamyloid (BB), sometimes slightly hemiamyloid (rB, very dirty reddish-grey at high concentration), of Sclerotinia-type, all parts equally reactive or sometimes upper ring paler, upper and lower ring deep blue when KOH-pretreated; base with (very) short and thick stalk arising from croziers {3}. Ascospores *(9.5–)11–13(–14.5) × (5–)5.5–6.5(–6.7) µm {5}, †(8.5–)9–11(–12.8) × 4.7–6.2(–6.5) µm {5}, ellipsoid to fusoid (homopolar) or slightly ovoid (heteropolar), ends rounded to obtuse, rarely subacute, ± equilateral; containing two medium-sized LBs (1.4–)1.8–2.4(–2.7) µm diam. {4}, each surrounded by a few small LBs, OCI (2–)3, sometimes with two small glycogen regions staining dextrinoid (red-brown in IKI), with 1 nucleus of 2.8–3 µm diam. in centre {1}; surrounded by a very delicate sheath that slips off the spore after ejection, spore wall surface CRB–; overmature 0–1-septate (septum median, rarely strongly eccentrical), *10.5–12.5 × 5–6 µm, without or with only small LBs; forming germ tubes and/or phialides. Paraphyses apically uninflated to slightly clavate {4}, in some apothecia frequently ± strongly clavate to capitate or spathulate-lageniform to moniliform {1}, terminal cell *(32–)52–85(–93) × 4–5.5(–6.5) µm {2}, †38–55 × 2.5–4.2 µm {1}, *6–10 µm wide if inflated, lower cells *(10–)20–32(–49) × 2.7–3.5 µm {2}; branched only in lower part; terminal cell containing large non-refractive vacuoles and a few very low-refractive globose SCBs; inflated apices laterally covered by a thin, pale ochraceous exudate. Subhymenium light ochre-brown, 20–30 µm thick, of dense textura intricata. Medullary excipulum subhyaline, *indistinctly/†distinctly gelatinized, of loose to dense t. intricata, individual cells *(40–)75–115 × 7–16 µm, smooth or slightly rough by some granular, pale ochraceous exudate, at base of receptacle 270 µm thick, at margin 80 µm, of dense, horizontal t. porrecta, sharply delimited from ectal excipulum; in stipe of vertical t. porrecta; hyaline stromatic tissue inside seed only observed close to insertion of stipe. Ectal excipulum ± hyaline, 50–100 µm thick at lower flanks, of thin-walled († slightly gelatinized, common walls 1–2 µm thick), ± vertically or often horizontally oriented t. globulosa(-prismatica) from base of receptacle up to mid flanks, cells *(16–)20–35(–70) × (12–)15–25(–35) µm {H.B. 8687}, †15–23 × 12–18 µm {H.B. 5698}, or †30–45 × 15–28 µm {H.B. 8955}, at mid flanks 30–40 µm thick, with ochre-brown exudate, marginal cells *38–70 × 6–10 µm {1}, hyphoid but terminally ± strongly inflated; exterior covered by 1–2 layers of *5–10 µm wide hyphae, these sometimes vertically projecting as 1–2-celled, short-cylindrical, hyaline hairs (†10–22 × 3–5 µm); in stipe of subhyaline to pale ochraceous, more basally bright red-brown t. porrecta, cells *28–46 × 3–4 µm, at very base of dark red-brown t. angularis; hairs on stipe scattered to dense, appressed or projecting, subhyaline, †40–80(–150) × 3–5 µm, at base light brown, †9–10.5 µm wide, with 0.5–1.3 µm thick smooth wall, covered with scattered granules. Rhomboid crystals very scattered to abundant in or on ectal excipulum {3}, especially at margin, sometimes also in hymenium, abundant in medullary excipulum of basal part of stipe and in stromatic tissue within seed, here forming druses 15–20 µm diam. Amyloidity of tissue: subhymenium {2} and outer medullary excipulum {1} distinctly pale blue (but negative at high concentration), or entire tissue IKI–, even if KOH-pretreated {2}. – Anamorph: Cultural characteristics: In pure culture on MEA and MMN the ascospores did not germinate. Sori formed mainly on the funiculus; conidiogenous cells not observed; chlamydospores singly, only sometimes in pairs, pale greyish-brown under transmitted light, blackish-brown under reflected light, coarsly warted and with irregular ridges, individual cells †(8.2–)8.7–11(–11.8) × 7.9–10.8 µm (without ornamentation) {H.B. 10206}, wall 0.7–1 µm thick, ornamentation ~0.2–0.5 µm high; germinating by phialides 7–14 × 3–6.5 µm on which microconidia 2.5–3.5 µm diam. are formed in (basipetal) chains {Vánky 1982}. Microconidial synanamorph formed on ascospores germinating in senescent apothecia, producing conidia either terminally or rarely laterally, directly on spore wall (sessile), or from small pegs 0.5–1.5 × 0.5–1.3 µm {H.B. 8687} or lageniform phialides †2.5–5 × 1.5–3 µm {H.B. 5698}; phialoconidia globose to subglobose, exceptionally subangular, *2.8–3.6 × 2.6–3.4 µm {H.B. 8687}, †2.5–2.9 µm diam. {H.B. 5698}, surface smooth or very indistinctly rough, CRB–; containing a single eccentrical LB (0.8–)1.2–1.6(–1.8) µm diam. {H.B. 8687}.
Habitat: Teleomorph on fallen, present or previous year's, moderately stromatized seeds of Veronica hederifolia agg. {11/6}, seeds 2–2.5 mm diam., surface always rugose, light to bright grey-brown, distinctly blackened only at a small area around insertion of stipe. Anamorph: chlamydospores formed apparently on placentae and funiculi of non-stromatized seeds of Veronica hederifolia s.str. {2/1} inside the initially closed capsule of living plants. Desiccation tolerance: apothecia dead in all parts after 1 day in the herbarium (except for ascospores); chlamydospores not tested. Phenology: apothecia: X–I, III–V; anamorph: V–VI (BubÁk 1916, SĂvulescu 1957, Zogg 1985, present collections); Altitude: 60–530 m. Geology: on basophilic to calcareous or more acidic soil: Muschelkalk (partly covered by loess), Lettenkeuper (clay), pleisto- & holocene sand & gravel and fluviatile sediments.
Phenology of Schroeteria decaisneana
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Specimens included (all on Veronica hederifolia agg.).
Teleomorph (on fallen seeds): Switzerland: Schaffhausen, 7 km NE of Schaffhausen, 1 km ENE of Thayngen, Flüheweg, MTB 8218/1, 530 m, ~10.I.1986, P. Blank & H.O. Baral (H.B. 2981). – Germany: Brandenburg, 25 km NW of Brandenburg, Rathenow, graveyard, 40–50 m, 29.X.1899, W. Kirschstein, vid. V. Kummer (B 70 0100003). – 3 km NE of Luckenwalde, WSW of Woltersdorf, Bürgerbusch, MTB 3845/3, 60 m, 9.V.1999, D. Benkert & V. Kummer, vid. V. Kummer (D.B., V.K., as Ciboria ploettneriana). – Sachsen-Anhalt, 5.5 km NE of Merseburg, NE of Burgliebenau, Elster-Luppe-Aue, MTB 4638/1, 85 m, 8.III.2009, G. Hensel (G.H. 080309). – Saale-Unstrutgebiet 4.3 km NE of Freyburg, 0.8 km W of Zeuchfeld, above vineyard, MTB 4736/42, 195 m, Muschelkalk (covered by Löss), 14.XI.2007, H. & U. Richter (U.R., sin. doc.). – ibid., 24.XI.2007, H. & U. Richter, P. & S. Rönsch (P.R., ex H.B. 8687, M-0276405). – ibid., 16.XI.2008, P. & S. Rönsch, U. Richter (P.R., H.B. 8955). – ibid., 29.XI.2009, U. Richter (U.R., A.U. 2273, sin. doc., sq.: ITS/LSU MZ048345). – ibid., 10.XII.2009, U. Richter (U.R., A.U. 2274, WU 43982, sin. doc., sq.: ITS/LSU MZ048346). – Bayern, Unterfranken, 7 km SSW of Schweinfurt, 1.5 km S of Grafenrheinfeld, near NSG Oberes Holz, Alter Main, MTB 6027/1, 200 m, 26.IV.1994, L.G. Krieglsteiner (L.K., H.B. 5698).
Anamorph (on seeds in capsules of living plants of V. hederifolia s.str.): Sachsen-Anhalt, 4.3 km NE of Freyburg, 0.8 km W of Zeuchfeld, above vineyard, MTB 4736/42, 195 m, 13.V.2019, H. & U. Richter (U.R., V.K. P1656/10, sin. doc.). – ibid., 21.V.2019, H. & U. Richter (H.B. 10206, J.K. S1346, GLM-F129032, sq.: ITS/LSU MW915644).
Taxonomic remarks: The first collection of the teleomorph of Schroeteria decaisneana was made in 1986 by P. Blank near Schaffhausen (Switzerland) and studied by the first author in the fresh state (Pl. 8: 1). The fungus was briefly reported by Baral (1986) as “Ciboria” cf. gemmincola Rehm, because it was erroneously believed to grow on galls of Neuroterus albipes (Cynipidae, Hymenoptera), a gall wasp that inhabits leaves of Quercus in Europe. In fact, oak galls, especially those of the related N. numismalis, resemble seeds of Veronica. In his review of Sclerotiniaceae on oak galls, Palmer (1991) mentioned the above collection by reproducing the first author's drawing, though without personal study. Ciboria gemmincola was described in Wagner (1895) on galls of Cynips gemmae with distinctly smaller, especially narrower spores of 8–9 × 3.5 µm.
In the following years the species was repeatedly collected in Sachsen-Anhalt (Pl. 7). At this occasion the fungus turned out to grow in fact on seeds of Veronica hederifolia agg. Therefore, the provisional name "Ciboria seminis-veronicae" was used for it, which appears also on the herbarium label of the specimen deposited in M as the intended holotype (H.B. 8687).
Variation: In all collections of Schroeteria decaisneana, from which detailed documentation was made, the paraphyses were cylindrical and apically scarcely inflated. Yet, in one of the large apothecia of a sample from Freyburg (Zeuchfeld, H.B. 8687) they were predominantly strongly inflated and variously shaped (Pls 7, 8: 3a, 11: 1j). The pale blue IKI-reaction of the subhymenium and outer medullary excipulum appears to be variable: it was present in the samples from Schweinfurt and Schaffhausen, but absent in those from Freyburg (no data available for the sample from Merseburg). Variation was also noted in ectal excipular cell size at the lower flanks, being much larger in the samples from Freyburg (H.B. 8687, 8955) compared to Schweinfurt (H.B. 5698) (no data available for those from Schaffhausen and Merseburg). Also the orientation of the cells varied. Finally, crystals were present in the collections from Schaffhausen and Freyburg, but they were not seen in those from Schweinfurt and Merseburg. In the Schweinfurt collection the germinating ascospores were predominantly 1-septate but also non-septate, and often they budded at both ends (Pl. 8: 2g), whereas in that from Freyburg they were always non-septate and germinated only at one conidiogenous locus (Pl. 8: 3c).
Cultural studies: Microconidia were abundantly produced from ascospores in senescent apothecia after incubation in a moist chamber. When shot on agar medium (MEA), the ascospores did not produce a mycelium even after several weeks of incubation at room temperature. According to studies by Brefeld (1912) and Vánky (1982), fresh chlamydospores germinated in tap water or strongly diluted nutrient solution at room temperature after 1–2 weeks and formed hyphae that abundantly produced phialides and microconidia. After adding concentrated nutrient solution, abundant mycelium developed instead. No attempts have been undertaken in the present study to obtain apothecia from stromatized seeds in a moist box.
Similar species: The study by the first author of a recent collection of Ciboria polygoni-vivipari Eckblad [Sweden, Lapland, Saxnäs, Marsfjället, 950 m, on bulbils of Bistorta vivipara (= Polygonum viviparum), 28.VII.2010, P. Perz, H.B. 9387, Baral ined., see IVV and https://svampe.databasen.org/taxon/11632] revealed some resemblance with the S. decaisneana teleomorph, particularly regarding the ascospores containing mostly two polar, medium-sized LBs. However, the spores were distinctly larger (*15–19 × 7.2–7.8 µm) and contained at least two nuclei, the paraphyses contained many low-refractive VBs in their apex, the ectal excipulum was of textura globulosa, and no crystals were seen. Ciboria seminicola (Kienholz & E.K. Cash) Hechler [including the questionable C. betulae (Woronin) W.L. White] on fruits of Alnus and Betula differs in somewhat longer and narrower, fusoid, warted ascospores and in asci arising from simple septa without croziers (Baral ined., H.B. 3677, 3682, 5136, 9774, see IVV).
Ecology: Schroeteria decaisneana was recorded near Freyburg (Zeuchfeld, Pl. 12) in a thermophilous deciduous forest with Ulmus minor bordered by Prunus spinosa, near Schweinfurt (Alter Main) in a riverbank forest (Querco-Ulmetum), near Merseburg (Elster-Luppe-Aue) in a riverbank forest with Prunus padus (?Pruno-Fraxinetum), and near Luckenwalde (Bürgerbusch) in a Pruno-Fraxinetum.The distribution of the anamorph of S. decaisneana comprises various countries of Europe (Scholz & Scholz 1988): Austria, former Czechoslovakia, France, Germany, Greece, Hungary, Italy, Poland, Romania, Sweden, Switzerland, and former Yugoslavia.
Schroeteria delastrina (Tul. & C. Tul.) G. Winter, Rabenh. Krypt.-Fl., Edn. 2 (Leipzig) 1.1: 117 (1881) [1884] – Pl. 15
≡ Geminella delastrina (Tul. & C. Tul.) J. Schröt., in Rabenh. Fungi Eur. exs. Cent. 14: no. 1376 (1870)
≡ Schizonella delastrina (Tul. & C. Tul.) Thirum. & M.D. Whitehead, Am. J. Bot. 55: 186 (1968)
≡ Thecaphora delastrina Tul. & C. Tul., Annls Sci. Nat., Bot., sér. 3 7: 108 (1847)
= Schroeteria delastrina var. reticulata Cocc., Mém. R. Accad. Sci. Ist. Bologna, Ser. 5 7: 219 (1898)
For a description see Vánky (1982).
Taxonomic remarks: A microscopic documentation of Schroeteria delastrina was done on the sample from Hannover (Pl. 15: 4c). Spore size was here evaluated as 7.8–11 × 7–9 µm (without ornamentation, see Tab. 4), the ornament as ~0.5–1(–2) µm high. According to Zogg (1985: 89, pl. 20B figs 1–5) the spore ornament is very different between S. delastrina and S. decaisneana, with warts and ridges up to 2 µm high in the former compared to only ~0.2 µm in the latter species.
Ecology: Schroeteria delastrina was reported by earlier workers on different Veronica species, though mainly on Veronica arvensis, the type substrate. Our investigated collections from Germany and Greece were all on this host plant which occurred at the collection sites in different biotops: in Germany in ruderal vegetation at the border of farmland (Kelbra), in mown meadows (Hannover, Thale), and in a Mesobromion grassland (Bad Frankenhausen, Leistadt, Sondershausen), in Greece in an annual vegetation on a raised area on the border of a summer-dry stream bed. The geology was mainly basophilically influenced, the phenology is May–June, in southern Europe March.
The distribution of the anamorph of S. delastrina comprises various countries of Europe (Scholz & Scholz 1988): Austria, Belgium, former Czechoslovakia, Denmark, Finland, France, Germany, Great Britain, Greece, Italy, Netherlands, Portugal, Poland, Romania, Sweden, Switzerland, former Soviet Union, and former Yugoslavia.
Specimens included (all in capsules of living plants of Veronica arvensis).
Anamorph: Germany: Thüringen, Kyffhäuser, 15 km SE of Nordhausen, 4 km WSW of Kelbra, between Mittelberg and Schloßberg Kleiner Heuweg, MTB 4531/44, 161 m, 28.V.2015, V. Kummer (V.K. P1652-23, sin. doc., sq.: ITS MW915652). – 0.9 km N of Bad Frankenhausen, Schlachtberg, 235 m, MTB 4632/23, 14.VI.2019, V. Kummer (V.K. P1652-26, sq.: ITS/LSU MW915645). – Hainleite, ESE of Sondershausen, N of Jecha, Panzerstraße 1, 185 m, MTB 4631/14, 14.VI.2019, V. Kummer (V.K. P1652-27, sin. doc., sq.: ITS/LSU MW915646). – Niedersachsen, 5.5 km ENE of Hannover, Groß Buchholz, Roderbruchmarkt, Nobelring, in front of hostel, MTB 3624/22, 55 m, 30.V.2011, J. Kruse (ex J. K. S0045, GLM-F129676). – Rheinland-Pfalz, N of Bad Dürkheim, Leistadt, 230 m, 13.V.2020, J. Kruse (J.K. S0969). – Hessen, Frankfurt/Main, SW airport, nearby Mönchhofdreieck, Flugschneise, 100 m, 24.V.2014, J. Kruse (J. K. S0239). – Darmstadt, Winkelschneise, forest sports park, ruderal area, 11.VI.2016, J. Kruse (J.K. S0604). – ~5 km SW of Darmstadt, L3097, Pfungstädter Hausschneise, 105 m, 11.VI.2016, J. Kruse (J.K. S0597) – 11 km NW of Eschwege, Frankershausen, Kripp- und Hielöcher, 270 m, 13.VI.2015, J. Kruse (J.K. S0335). – Nordrhein-Westfalen, Essen-Frohnhausen, Martin-Luther-Straße, 75 m, 31.V.2017, J. Kruse (J. K. S0956) – Sachsen-Anhalt, 11 km SW of Quedlinburg, Thale, Hubertusstraße, “Friedenspark”, 180 m, 9.VI.2017, J. Kruse (J.K. S0969). – L208 between Balgstädt and Hirschroda, 2,1 km WNW of Balgstädt, Balgstädter Berge, 24.V.2017, J. Kruse (J.K. S0953). – Berlin, 15 km SE of Berlin, Altglienicke, 50 m, 17.V.2014, V. Kummer (ø). – Bayern, ca. 2,4 km W Ibind, Fitzendorfer Wollgraswiesen, 340 m, 20.VI.2020, J. Kruse (ø). – Coburg, ~1 km NNW of Gemünda, Heiligenwiesen/Heiligenleite, 310 m, 21.VI.2020, J. Kruse (ø). – Greece: Rhodos, 4 km SW of Malonas, 1.8 km SSE of monastery Kamiri, entrance to zur Scoutljaris gorge, 60 m, 29.III.2011, V. Kummer (V.K. P1652-20).
Schroeteria bornmuelleri Magnus, Mitt. Thür. Bot. Ver., N.F. 28: 64 (1911)
For a description see Vánky (1982).
Specimens included (in capsules of living plants of Veronica rubrifolia).
Anamorph: Iran: Khorasan, 40 km NE of Mahhad, Gughghi (Gojgi), ~1250 m, 11.V.1990, D. Ershad, T. & K. Vánky (H.U.V. 750 ex TUB, sq.: ITS MW915653; BRIP H.U.V. 14892, 15006, non. vid.).
Schroeteria poeltii Vánky, Mycotaxon 18(2): 326 (1983) — Pls 16–19
Etymology: named after the Austrian lichenologist Josef Poelt (1924–1995) who sent the Hungarian smut fungi specialist Kálmán Vánky (*1930) some samples of the fungus.
Holotype: France, Alpes-Maritimes, Menton, Ste. Agnés, seeds of Veronica cymbalaria, 20.VI.1962, H. Teppner (BRIP H.U.V. 10800, non. vid.).
DESCRIPTION. Teleomorph: Apothecia fresh (1–)1.3–2.5(–3) mm diam., non-gelatinous, disc light ochraceous, slightly concave, finally flat, margin thin, not protruding, ± smooth, exterior pale ochraceous, very finely pubescent, receptacle at base 0.5–0.75 mm thick, at margin 0.25–0.35 mm; stipe 5–17 × (0.07–)0.15–0.55 mm, concolorous with receptacle in upper part, darker red-brown in lower part, black-brown at base, overall densely pubescent with rather long hairs; often gradually thinner towards base, flexuous especially in lower part, emerging singly from convex or lateral side of the seed. Asci *145–160 × 9.5–10.5(–10.8) µm {1}, †114–135 × 7–8.5 µm {1}, protruding *5–30 µm beyond paraphyses or (†) ± equalling the paraphyses, with 8 equal-sized spores (in overmature apothecia also unequal-sized), pars sporifera *53–63 µm long, spores obliquely subbiseriate; apex (†) hemispherical to moderately truncate, apical thickening immature or mature †2–3.8 µm thick, apical ring blue in IKI {2}, euamyloid (BB), of Sclerotinia-type, lower ring strongly blue, upper parts less so; base with short and thick stalk, arising from croziers {1}. Ascospores *(9.5–)10–11(–11.3) × (5.3–)5.5–6(–6.3) µm {1}, †(8.6–)8.8–10(–10.3) × 4.7–5(–5.3) µm {1}, ellipsoid (homopolar) or very slightly ovoid (heteropolar), ends rounded to obtuse, ± equilateral; containing 1(–2) medium-sized LBs close to each end, (1–)1.5–1.8 µm diam. {3}, OCI 2, glycogen not observed, with 1 nucleus 3–3.2 µm diam. in centre {1}; without sheath; overmature spores not observed. Paraphyses apically uninflated to slightly clavate-capitate or sublageniform {1}, terminal cell *57–70 × 3.5–4.7(–5.2) µm {1}, †3–4.1 µm wide, projecting beyond †asci by 5–10 µm; terminal cell containing large non-refractive vacuoles. Subhymenium light ochre-brown, 25–35 µm thick, of dense textura intricata. Medullary excipulum subhyaline, non-gelatinized, of dense horizontal t. intricata-porrecta, individual cells *(40–)70–130 × 6–11 µm, smooth or slightly rough by some granular, pale ochraceous exudate, 100–130 µm thick at lower flanks, sharply delimited from ectal excipulum; in stipe of vertical t. porrecta, cells *60–90 × 7–13 µm. Ectal excipulum 80–90 µm thick at lower flanks, ± hyaline, at mid flanks 25–35 µm thick, brown; of thin-walled, ± vertically oriented t. globulosa(-prismatica) from base of receptacle up to mid flanks, cells *20–50 × 15–30 µm {1}; exterior without covering hyphae but with scattered to abundant, protruding, vesiculous or cylindrical to conical, hyaline to brownish, 0–1-septate hair-like cells or hairs *25–35 × 13–15 µm, †18–25 × 3–10 µm; in stipe of subhyaline to pale ochraceous t. prismatica-porrecta, cells †25–40 × 6–9.5 µm; hairs on stipe scattered to dense, appressed or projecting, subhyaline to pale brown, †(22–)35–195(–258) × (3–)4–6(–8) µm, with 0.3–0.7 µm thick smooth wall, covered with scattered granules, sparsely septate, individual cells ~38–85(–140) µm long, usually emerging from superficial, swollen basal cells †15–32 × 11–17 µm. Rhomboid crystals scattered in medullary and ectal excipulum, more abundant in stipe. Amyloidity of tissue: subhymenium weakly pale blue (KOH-pretreated) {1}. – Anamorph: Cultural characteristics: ascospores germination not examined. Sori forming a reddish-brown mass of chlamydospores under reflected light which completely replace the seeds; conidiogenous cells not observed; chlamydospores cohering to form strongly curved (U-shaped) chains of (2–)4–5(–7) spores {2}, light yellowish-brown under transmitted light, ± smooth or often with distinct low warts, individual cells *10–11 × 8–10 µm {1}, †(5.5–)6.5–12 × 6.5–13 µm {type}, multiguttulate, wall 0.5–0.8 µm thick, ornamentation ~0.3–1(–2) µm high. Microconidial synanamorph (formed on chlamydospores, evaluated from Nagler et al. 1989) hyphoid conidiophores with conidiogenous cells phialidic, cylindrical to lageniform, 8–20 × 3.5–5.8 µm, microconidia ± globose, *(2.7–)3–3.7(–4) × (2.7–)2.9–3.5(–3.8) µm, with one eccentrical LB of 1.5–2 µm diam.
Habitat: Teleomorph on fallen, previous year's, moderately stromatized seeds of Veronica cymbalaria {1}, seeds (0.9–)1.7–2.8 mm diam., surface always rugose, light to bright grey-brown or blackish. Anamorph: chlamydospores formed apparently on placentae and funiculi of non-stromatized capsules of V. cymbalaria {3/3} by replacing the seeds inside the initially closed capsule of living plants. Desiccation tolerance: not tested. Phenology: apothecia I; anamorph III, VI. Altitude: 7–650 m. Geology: on acidic and calcareous soil.
Specimens included (all on Veronica cymbalaria).
Teleomorph (on fallen seeds): Spain: Andalucía, Málaga, 14 km S of Ronda, N of Pujerra, arroyo Bollage, 590 m, 4.I.2017, F.J. Valencia, vid. R. Tena (ex C.V.L. 040117, JA-CUSSTA 9523, GLM-F29000, sq.: ITS/LSU MW915648).
Anamorph (on seeds in capsules of living plants): France: Provence-Alpes-Côte-d'Azur, Alpes-Maritimes, ~4 km NW of Menton, near Ste.-Agnès, ~600 m, 20.VI.1962, H. Teppner (BRIP H.U.V. 10800, holotype, GZU 292883 & 294859 isotypes, doc. vid.). – ibid., ~650 m, 10.VI.1987, K. Vánky (BRIP H.U.V. 13121, topotype, doc. vid.). – Greece: Rhodos, 1.7 km SSW of Ialysos, 0.2 km E of Filerimos, monastery park, 212 m, 20.III.2018, J. Kruse & V. Kummer (V.K. P1675/cymbalaria 1, J.K. S1304, sq.:.ITS MW915654). – 2.7 km SSE of Masari, SW of Vagies, coastal road, 5 m, 22.III.2018, J. Kruse (J.K. B2278, sq.: ITS/LSU MW915647) – 2.5 km SE of Masari, Charaki, coastal road, former barracks area on the beach, 0 m, 21.III.2018, J. Kruse (J.K. B2279).
Taxonomic remarks: In the teleomorph, Schroeteria poeltii deviates from S. decaisneana merely by slightly smaller asci and ascospores, the latter having their LBs a bit closer to the spore ends, perhaps also in the absence of glycogen in the ascospores and in a tendency to wider excipular cells (Tab. 3). Despite these minor differences, the anamorph sharply differs by smooth, yellowish-brown chlamydospores cohering in strongly curved (U-shaped) chains of (2–)4–7 spores from the other Schroeteria spp. with their warted, greyish-brown chlamydospores cohering in straight chaines of 2–4 spores or detaching as single spores. The exclusive occurrence of S. poeltii on V. cymbalaria and the remarkable anamorph make any confusion with other species very unlikely.
Nagler et al. (1989) investigated the type of Schroeteria poeltii and a collection of S. delastrina on V. arvensis by light and electron microscopical methods. The authors observed microconidia formed endogenously in phialides which emerge either from germ tubes of the chlamydospores or directly from the chlamydospores. Besides, the authors illustrated an unusual case of endogenous maturation of microconidia inside of chlamydospores (fig. 7), which they also observed inside hyphal cells of S. delastrina (fig. 9). Problematic is that the scales are wrong in some of their illustrations. In order to achieve reasonable measurements, the scales in figs 6 and 9 should be around 20 µm instead of 5 µm and that in fig. 7 around 3 µm instead of 2 µm, whereas the scales in figs 12–17 appear to be correct.
In the collection from Filerimos, not all capsules of a given plant individuum were filled with chlamydospores but numerous were with seed formation. However, this varied even on a single shoot of a plant. Whether the seeds are able to germinate has not been tested. This is in contrast to our collections of Schroeteria delastrina on Veronica arvensis where all capsules of a shoot on a plant were either filled with chlamydospores or with seeds, suggesting a systemic infection.
Typification and etymology: Vánky (1983), who investigated the type of S. poeltii, stated that the species was only known from the type locality, where it was collected by H. Teppner in 1962. In a footnote he explained the reason for naming the species S. poeltii: shortly after publication of Vánky's Schroeteria monograph which appeared in 1982, Josef Poelt had sent him a specimen of Teppner's collection. Part of this collection then remained in GZU, whereas the holotype was stated by Vánky to have been deposited at UPS. However, according to Å. Kruys (pers. comm.) the specimen could not be found at UPS. In 1987 Vánky collected a topotype, which Nagler et al. (1989) investigated. In 2013 Vánky gave his entire herbarium to BRIP (Brisbane) where the holotype and also the topotype are listed in the online database.
Ecology: The host plant of Schroeteria poeltii has a (sub)mediterranean distribution and forms urn-shaped seeds similar as in V. hederifolia (cyathiform fide Muñoz-Centeno et al. 2006).
The French holotype collection was from a mesomediterranean semihumid site northeast of Monaco in dépt. Alpes-Maritimes. The collecting locality is not fully clear: the rough coordinates (43° 47' N, 7° 30' E) given by Vánky (1983) are in a region about 1 km north of the seaside town Menton and cover a region of about 50–200 m altitude. The given data about altitude (ca. 600 m) and site (tract Ste. Agnès, Umgebung von Ste. Agnès) in Vánky (l.c.) and on the handwritten isotype label in GZU (which lacks coordinates), suggest that the collection site was about 4 km northwest of Menton, i.e., about 3 km away from the published coordinates.
The Schroeteria poeltii anamorphs from thermomediterranean semihumid Greece were collected at the site Filerimos in an open grove with Quercus coccifera on top of a hill in the monastery park at 212 m altitude in the north of Rhodos, and near Charaki in a ruderal farmland area with Mercurialis annua etc., a former military area close to coastline at 7 m altitude in the middle east of Rhodos. The teleomorph from supramediterranean semihumid Spain was from an acidic floodplain forest in Málaga (Andalucía). The apothecia were found on stromatized seeds of Veronica cymbalaria fallen to the ground. The plants grew just above the ground on the vertical area of a schist rock. The floodplain forest consisted of Populus alba and Salix alba, and mediterranean plants such as Quercus faginea and Rubia peregrina, also Lamium flexuosum, Rubus ulmifolius, Dorycnium rectum, Ranunculus ficaria, and Vinca difformis. Outside of the flooded area are large Castanea sativa plantations, the main agricultural product of the local people.