A new approach to species delimitation in Septoria - CBS - KNAW

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Verkley et al.to 60 µm diam, lacking distinctly differentiated cells; conidiomatalwall composed of textura angularis without distinctly differentiatedlayers, mostly 15–20 µm thick, the outer cells with brown,somewhat thickened walls and 4.5–10 µm diam, the inner cellshyaline and thin-walled and of comparable diam. Conidiogenouscells hyaline, discrete or integrated in short, 1–2-septateconidiophores, cylindrical, or ampuliform with a relatively shortneck, hyaline, holoblastic, proliferating sympodially, and sometimesalso percurrently with indistinct annellations, 6.5–10(–12.5) × 2.5–4.5 µm. Conidia cylindrical, weakly to strongly curved, or flexuous,gradually attenuated to a rounded apex, gradually or more abruptlyattenuated into a broadly truncate base, (0–)2–5(–6)-septate, notor indistinctly constricted around the septa, hyaline, contents withseveral small guttules and numerous granules in each cell in theliving state, oil-droplets rarely merged into larger guttules in therehydrated state, (20–)40–65 × 2–2.5(–3) µm (rehydrated). Sexualmorph unknown.Description in vitro: Colonies on OA 7–10 mm diam in 2 wk (22–26 mm in 6 wk), with an even, somewhat undulating, glabrous,colourless margin; colonies spreading, the surface plane, immersedmycelium pale luteous or buff, with scattered immersed andsuperficial pycnidial conidiomata, which are first dark olivaceous,then almost black, glabrous, 150–450 µm diam, with a single orseveral (up to 5!) ostioli placed on short papillae or more elongatednecks (up to 350 µm), that release buff to rosy-buff later salmonconidial slime; aerial mycelium diffuse, woolly-floccose, white;reverse honey, but isabelline to hazel in the centre. Colonies onCMA 6–8 mm diam in 2 wk (18–23 mm in 6 wk), with an even,glabrous margin; as on OA but immersed mycelium with a greenishhaze; aerial mycelium higher and reverse darker, later hazel witholivaceous and yellow tinges; conidiomata similar as on OA.Colonies on MEA 7–9 mm diam in 2 wk (18–21 mm in 6 wk), withan even or somewhat undulating, glabrous, buff to honey margin;colonies pustulate to almost hemispherical, immersed myceliumrather dark, near the margin covered by woolly to felty white aerialmycelium; mostly composed of spherical conidiomatal initials,superficial mature conidiomata releasing rosy-buff to salmon, laterhoney conidial slime; reverse dark brick in the centre, near themargin cinnamon to honey. Colonies on CHA 7–14 mm diam in 2wk (20–28 mm in 6 wk), with an irregular, buff margin covered by adiffuse, felty white, later grey aerial mycelium; further as on MEA,but the colony surface less elevated and especially near the marginwith greyish felty to tufty aerial mycelium; conidial slime abundantlyproduced, first rosy-buff, later salmon to ochreous; reverse in thecentre blood colour, dark brick to cinnamon at the margin.Conidiomata on OA see above. Conidia as in planta, mostly(0–)3–5(–6)-septate, 44–63(–70) × 2.5–3 µm.Hosts: Senecio fluviatilis and S. nemorensis.Material examined: Belgium, Château de Namur, on leaves of Senecio sarracenica,1829, A. Bellynck, isotype BR-MYCO 155500-09. Netherlands, Prov. Gelderland,Millingen a/d Rijn, Millingerwaard, on living leaves of S. fluviatilis, 6 Oct. 1999, G.Verkley 939, epitype designated here CBS H-21219 “MBT175358”, living culturesex-epitype CBS 102366, 102381.Notes: The first Septoria that was described on the genus Seneciowas S. senecionis. The type host is Senecio sarracenica (=Senecio fluviatilis), and in later literature it has also been reportedfrom several other species of Senecio (Radulescu et al. 1973).According to the diagnosis by Westendorp, the conidia are 40 ×1.5 µm and 3–4-septate. Vanev et al. (1997) described the conidiaof S. senecionis as 2–6-septate, 29–68 × 2–2.5 µm, Radulescuet al. (1973) as 3–4-septate, 33–57 × 1.2–2 µm. By examiningthe type specimen from BR it is here confirmed that conidia are infact wider than described by Westendorp. It contains a single leafwith a few lesions, and conidia observed are 30–55 × 1.5–2.5 µm,and mostly 3–5-septate. The fresh material that was collected inthe Netherlands from the same host species, Senecio fluviatilis,and from which CBS 102366 and 102381 were isolated, is insufficient agreement with the type and is therefore designated hereas epitype of S. senecionis. Differences with Septoria putrida arediscussed under that species.Septoria sii Roberge ex Desm., Pl. crypt. Fr., Fasc. 44, no2185; Annls Sci. Nat., sér. 3, Bot. 20: 92. 1853. Fig. 37.Description in planta: Symptoms leaf spots, yellow to brown, initiallyvaguely delimited but later well-delimited by veinlets, scattered,later often confluent over large areas, visible on both sides of theleaf. Conidiomata pycnidial, epiphyllous, rarely also hypophyllous,single, scattered or in small clusters, globose to subglobose,immersed, (60–)80–110 µm diam; ostiolum circular, central, 12.5–25(–35) µm wide, surrounding cells concolorous; conidiomatal wallcomposed of textura angularis 5–10 µm thick, with an outer layer ofcells 3–4.5 µm diam with brown, thickened walls, and an inner layerof hyaline and thin-walled cells, 2.5–4 µm diam. Conidiogenouscells hyaline, broadly or elongated ampulliform, normally witha distinct neck, hyaline, holoblastic, proliferating percurrently,annellations indistinct, 5–8.5 × 3–5 µm. Conidia cylindrical, straight,curved, or flexuous, gradually attenuated to a relatively broadlyrounded apex, more or less abruptly attenuated into a truncatebase, 1–3(–4)-septate, slightly to distinctly constricted around thesepta in the fresh, fully hydrated state, hyaline, containing one toseveral relatively large oil-droplets in each cell, in the rehydratedstate with irregular oil-masses (20–)29–35(–42) × 2–2.5(–3) µm(living; rehydrated, 1.5–2 µm wide). Sexual morph unknown.Description in vitro: Colonies on OA 4–9 mm diam in 2 wk [(15–)19–23 mm in 6 wk], with an even, glabrous, colourless margin;colonies remaining almost plane, immersed mycelium olivaceousblack,locally however peach is dominant, which becomes scarletafter several wk; aerial mycelium mostly well-developed, woollyfloccose,white; scattered, mostly immersed pycnidial to stromaticconidiomata developing in the centre, releasing droplets ofmilky white to rosy-buff conidial slime; reverse dark slate blue toolivaceous-black, and locally peach, the pigment not diffusing intothe medium. Colonies on CMA up to 1.5 mm diam in 2 wk [7–10(–25) mm in 6 wk], as on OA, but peach pigment diffusing into themedium, while the colony itself is predominantly olivaceous-black.Colonies frequently develop faster growing sectors that first arebuff and sporulate directly from the mycelium, later become paleluteous with a distinct scarlet pigmentation and forming numerousmostly superficial pycnidia. Colonies on MEA 3–6 mm diam in2 wk [12–14(–26) mm in 6 wk], the margin ruffled, olivaceousblack;colony concolorous, irregularly pustulate-worty, covered bydiffuse to dense felty white or greyish aerial mycelium; numerousconidiomatal initials developing at the surface, mature onesreleasing cirrhi of conidia that first are milky white, later salmon,sometimes merging to form slimy masses covering areas of thecolony surface; the agar surrounding the colony slightly discolouredby diffusing pigment(s). Colonies on CHA 5–6 mm diam in 2 wk[8–13(–15) mm in 6 wk], as on MEA; some parts of the colonies286
A new approach to species delimitation in SeptoriaFig. 37. Septoria sii. A–C. Colonies CBS 102370 (15 °C, nUV). A. On OA. B. On CHA. C. On MEA. D. Conidia and conidiogenous cells in planta (CBS H-21223). E. Ibid., onOA (CBS 102369). Scale bars = 10 µm.pale ochreous, tardily sporulating, releasing pale flesh to salmondroplets of conidial slime from superficial pycnidial conidiomata.Cultures sporulating with conidiogenous cells developing in(superficial) mycelial hyphae, solitary or in sequences, in additionto conidiomata. Conidiomata on OA pycnidial, single, dark brownto black, 80–185 µm diam, ostiolum single 30–60 µm diam, orstromatic without a differentiated opening and up to 220 µm diam;conidiogenous cells inside pycnidia as in planta but often withmore elongated neck, holoblastic, percurrently proliferating one toseveral times with indistinct annellations, 7–12.5 × 3–6 µm. Conidiaas in planta, 22–43 × 2.2–2.5 µm.Hosts: Sium latifolium, other Sium spp. and Berula erecta (syn.Sium erectum).Material examined: Netherlands, Prov. Friesland, Terschelling, ditch in polder Sof Hoorn, on living leaves of Berula erecta, 19 Aug. 1995, H.A. van der Aa 12029,CBS H-18173, living culture CBS 118.96; same substr., Prov. Utrecht, ‘s Graveland,Kortenhoefse plassen, “Oppad”, 14 Oct. 1999, G. Verkley & H.A. van der Aa 945,CBS H-21223, living culture CBS 102369; same loc., substr., date, G. Verkley & H.A.van der Aa 946, CBS H-21222, living culture CBS 102370.Notes: The stout conidia with blunt apices and distinct constrictionsaround the septa (at least in the living, turgescent state) and theabsence of sympodial proliferation in conidiogenesis distinguish thisspecies from most other Septoria on Apiaceae here investigated,including S. apiicola. According to the original diagnosis, based onmaterial from Sium latifolium in France, the conidia are 30–40 ×2.5 µm. Most later authors have reported somewhat different sizeranges; for example Teterevnikova-Babayan (1985) observedconidia 20–60 × 1–1.5 µm, Vanev et al. (1997) 20–41 × 1.5–2.2 µm,and Radulescu et al. (1973) reported 30–40 × 2–3 µm. The materialavailable for this study proved homogeneous in morphology andgenotype. The phylogenetic data indicate that this species is veryclosely related to S. mazi, a fungus occurring on Mazus japonica(Scrophulariaceae), but also to S. aegopodina on Aegopodium sp.(Apiaceae). The conidia of S. mazi morphologically resemble thoseof S. sii, but they are narrower and the septa normally indistinct[15–42 × 1.5–2(–2.5) µm, Shin & Sameva 2004].Septoria sisyrinchii Speg., An. Mus. nac. Hist. nat. B. Aires,6: 324. 1899. Fig. 38.Description in planta: Symptoms leaf lesions developing in largeareas of the leaf lamina that first turn yellow, indefinite, hologenous,pale to dark brown, appearing black due to numerous conidiomata.Conidiomata pycnidial, amphigenous, numerous, semi-immersedto immersed, subglobose to globose, black, 70–100(–120) µmdiam; ostiolum central, circular, 15–35 µm wide, sometimesopening more widely, releasing white to pale yellowish cirrhi ofconidial slime, surrounding cells concolourous or somewhat darker;conidiomatal wall 15–20 µm thick, composed of an outer layer ofisodiametric cells 5–8 µm diam with brown, slightly thickened cellwalls up to 1 μm thick, and an inner layer of globose to isodiametriccells 3–6 μm diam with thin, hyaline walls. Conidiogenous cellshyaline, discrete or integrated in 1-septate conidiophores up 15µm long, cylindrical, or ampulliform, holoblastic, proliferatingsympodially, percurrent proliferations not observed, 5–10 × 2.5–3.5µm. Conidia cylindrical to cylindrical-filiform, slightly to stronglycurved, sometimes flexuous, narrowly rounded to somewhatpointed at the apex, attenuated gradually or more abruptly towardswww.studiesinmycology.org287
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Verkley et al.CaryophyllaceaeRutace
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Verkley et al.Fig. 3. Caryophyllose
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Page 54 and 55: Verkley et al.much more profound di
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Page 60 and 61: Verkley et al.to very dark dull gre
Page 62 and 63: Verkley et al.Fig. 30. Septoria nap
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Page 78 and 79: Verkley et al.on OA, but olivaceous
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