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

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

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Verkley et al.Sep<strong>to</strong>ria galeopsidis Westend., Bull. Acad. r. Belg., Cl. Sci.,Sér. 2, 2: 577. 1857. Fig. 20.= Ascochyta galeopsidis Lasch <strong>in</strong> Rabenh., Herb. Myc. I, 1058. 1846 [nom.nud.].= Sep<strong>to</strong>ria cotylea Pat. & Har., Bull. Soc. Mycol. France 21: 85. 1905.Description <strong>in</strong> planta: Symp<strong>to</strong>ms leaf spots irregular or angular,becom<strong>in</strong>g dark brown, <strong>in</strong> yellow parts of the leaf lam<strong>in</strong>a.Conidiomata pycnidial, hypophyllous, often numerous <strong>in</strong> eachleaf spot, globose <strong>to</strong> subglobose, dark brown, almost completelyimmersed, 75–100(–130) µm diam; ostiolum central, <strong>in</strong>itiallycircular, 15–25 µm wide, surround<strong>in</strong>g cells somewhat darker;conidiomatal wall 10–22 µm thick, composed of textura angulariswithout dist<strong>in</strong>ctly differentiated layers, the cells 3–8 µm diam,the outer cells with brown, somewhat thickened walls, the <strong>in</strong>nercells with hyal<strong>in</strong>e and th<strong>in</strong>ner walls. Conidiogenous cells discrete,sometimes <strong>in</strong>tegrated <strong>in</strong><strong>to</strong> 1–2-septate conidiophores, hyal<strong>in</strong>e,narrowly or broadly ampulliform with a relatively narrow neck,holoblastic, proliferat<strong>in</strong>g percurrently with <strong>in</strong>dist<strong>in</strong>ct annellations,and also sympodially, 6–12(–15) × 3.5–5(–6) µm. Conidia filiform,straight or slightly curved, sometimes flexuous, with a rounded orsomewhat po<strong>in</strong>ted apex, attenuated <strong>to</strong>wards the narrowly truncatebase, (0–)3(–5)-septate, not constricted around the septa, hyal<strong>in</strong>e,contents with several m<strong>in</strong>ute oil-droplets and granular material <strong>in</strong>each cell <strong>in</strong> the liv<strong>in</strong>g state, with <strong>in</strong>conspicuous oil-droplets andgranular contents <strong>in</strong> the rehydrated state, 20.5–44 × 1.5–2.5 µm(liv<strong>in</strong>g; rehydrated, 1–2 µm wide). Sexual morph unknown.Description <strong>in</strong> vitro: Colonies on OA 7–13 mm diam <strong>in</strong> 2 wk (35–43mm <strong>in</strong> 6 wk), with an even, glabrous, colourless marg<strong>in</strong>; coloniesalmost plane, immersed mycelium homogeneously olivaceousblack<strong>to</strong> greenish black (also near the marg<strong>in</strong>); aerial myceliumscanty, woolly-floccose, white or greyish; superficial pycnidialconidiomata scanty, scattered over the central aerea, releas<strong>in</strong>gmilky white droplets of conidial slime; reverse dark slate blue <strong>to</strong>black. Colonies on CMA 7–13 mm diam <strong>in</strong> 2 wk (33–37 mm <strong>in</strong> 6wk), as on OA, but concentration of conidiomatal development <strong>in</strong>elevated pustules on the elsewhere flat colony. Colonies on MEA6–11 mm diam <strong>in</strong> 2 wk (33–39(–46) mm <strong>in</strong> 6 wk), the marg<strong>in</strong> even,later undulat<strong>in</strong>g, buff, narrow and glabrous; colonies hemispherical,often irregularly pustulate or with columnar outgrowths up <strong>to</strong> 5mm high, immersed mycelium olivaceous-black <strong>to</strong> black, mostlycovered by a dense mat of f<strong>in</strong>ely velted, greyish aerial mycelium;faster grow<strong>in</strong>g, glabrous sec<strong>to</strong>rs with buff immersed mycelium mayappear after several weeks; conidiomata start<strong>in</strong>g <strong>to</strong> develop onthe (dark) colony surface, tardily sporulat<strong>in</strong>g with whitish <strong>to</strong> fleshdroplets of conidial slime; reverse brown-v<strong>in</strong>aceous or olivaceousblack.Colonies on CHA 5–10(–15) mm diam <strong>in</strong> 2 wk (20–29 mm<strong>in</strong> 6 wk), with an even, glabrous <strong>to</strong> nearly so, buff marg<strong>in</strong>; coloniesirregularly pustulate, immersed mycelium olivaceous-black, mostlycovered by a dense but appressed mat of woolly-floccose, greyaerial mycelium, <strong>in</strong> some slightly faster grow<strong>in</strong>g sec<strong>to</strong>rs pure white;scattered but scarce superficial conidiomata releas<strong>in</strong>g pale fleshdroplets of conidial slime; reverse blood colour <strong>to</strong> black.Conidiomata pycnidial and similar as <strong>in</strong> planta, 100–150 µmdiam, or merged <strong>in</strong><strong>to</strong> larger complexes especially on the agarsurface, dark brown, up <strong>to</strong> 200 µm diam; ostiolum as <strong>in</strong> planta,or absent. Conidiogenous cells hyal<strong>in</strong>e, ampuliform, or elongatedampulliform <strong>to</strong> cyl<strong>in</strong>drical, with a dist<strong>in</strong>ct neck, holoblastic,proliferat<strong>in</strong>g percurrently with <strong>in</strong>dist<strong>in</strong>ct scars (annellations), orsympodially, 8–13(–15) × 3–4.5(–5) µm. Conidia cyl<strong>in</strong>drical,straight or slightly curved, taper<strong>in</strong>g <strong>to</strong> a rounded or somewhatpo<strong>in</strong>ted apex, lower part slightly or more clearly attenuated <strong>in</strong><strong>to</strong> abroad truncate base, (0–)1–3(–5)-septate, not constricted aroundthe septa, hyal<strong>in</strong>e, with several oil-droplets and m<strong>in</strong>ute granularmaterial <strong>in</strong> each cell, (37–)50–65 (–70) × 2–2.5 µm.Hosts: Galeopsis angustifolia, G. ladanum, G. pubescens, G.speciosa and G. tetrahit.Material exam<strong>in</strong>ed: Belgium, <strong>in</strong> the vic<strong>in</strong>ity of Mons, on leaves of Galeopsis tetrahit,R. P. Clém. Dumont, distributed <strong>in</strong> Westendorp & Wallays, Herb. crypt. Belge, Fasc.23-24, no 1134, isotype BR-MYCO 158116-06. Czech Republic, Moravia, Mikulov,on liv<strong>in</strong>g leaves of Galeopsis sp., 15 Sep. 2008, G. Verkley 6003, <strong>CBS</strong> H-21256,liv<strong>in</strong>g cultures <strong>CBS</strong> 123744, 123749; same substr., date, Moravia, Milovice, forestMilovika stran, G. Verkley 6006, <strong>CBS</strong> H-21254, liv<strong>in</strong>g cultures <strong>CBS</strong> 123745, 123746.France, Corrèze, Prât Alleyrat, on liv<strong>in</strong>g leaves of G. tetrahit, 25 July 1976, H.A. vander Aa 5344, <strong>CBS</strong> H-18099; loc. unknown, isol. C. Killian ex Galeopsis sp., liv<strong>in</strong>gculture <strong>CBS</strong> 191.26. Netherlands, prov. Noord-Brabant, Cromvoirt, on liv<strong>in</strong>g leavesof G. tetrahit, 2 June 1963, H.A. van der Aa s.n., <strong>CBS</strong> H-18097; prov. Gelderland,Putten, on liv<strong>in</strong>g leaves of G. tetrahit, 8 Aug. 1984, G. de Hoog s.n., <strong>CBS</strong> H-18100;prov. Utrecht, Soest, on liv<strong>in</strong>g leaves of G. tetrahit, 4 Aug. 1999, G. Verkley 902, <strong>CBS</strong>H-21195, liv<strong>in</strong>g culture <strong>CBS</strong> 102314; prov. Limburg, St. Jansberg near Plasmolen,on liv<strong>in</strong>g leaves of G. tetrahit, 9 Sep. 1999, G. Verkley 934, epitype designatedhere <strong>CBS</strong> H-21215 “MBT175356”, liv<strong>in</strong>g culture ex-epitype <strong>CBS</strong> 102411. Romania,distr. Satu-Mare, Pir, on liv<strong>in</strong>g leaves of G. ladanum, 27 Aug. 1973, G. Negrean s.n.,<strong>CBS</strong> H-18098.Notes: Jørstad (1965) reported comparable conidial size ranges<strong>in</strong> specimens on different host <strong>species</strong>, viz. G. speciosa (extremevalues 20–64 × 1–2.5 µm) and G. tetrahit (28–60 × 1–2 µm),although <strong>in</strong> most Norwegian collections on G. tetrahit, the maximumconidial length varied downwards <strong>to</strong> 48 µm. In the orig<strong>in</strong>aldiagnosis of S. galeopsidis conidia are described as 30–40 ×1–1.5 µm (Saccardo 1884), while Radulescu et al. (1973) reportedmeasurements rang<strong>in</strong>g between 20–45 µm <strong>in</strong> length <strong>in</strong> collectionson various hosts. In the type material from BR <strong>in</strong>vestigated hereconidia are mostly 3–5-septate, 19–40 × 1.5–2 µm. In othermaterial available for the present study, maximum length of conidiawas only 44 µm <strong>in</strong> planta, whereas the stra<strong>in</strong>s obta<strong>in</strong>ed from it werecapable of form<strong>in</strong>g conidia with a maximum length of 70 µm on OA.The differences with S. lamiicola are discussed under that <strong>species</strong>.Sep<strong>to</strong>ria galeopsidis is closely related <strong>to</strong> only some of the otherSep<strong>to</strong>ria <strong>species</strong> occurr<strong>in</strong>g on plants from the family Lamiaceae,especially S. melissae (<strong>CBS</strong> 109097) and S. stachydis. Sep<strong>to</strong>rialamiicola on Lamium spp., which is morphologically quite similar<strong>to</strong> S. galeopsidis, proves genetically very dist<strong>in</strong>ct, although thesetaxa can barely be dist<strong>in</strong>guished by their ITS sequence (99.5 %).Several house-keep<strong>in</strong>g genes do allow an easy identification ofthese <strong>species</strong>.Sep<strong>to</strong>ria heraclei (Lib.) Desm., Pl. crypt. Fr., Fasc. 11, no534. 1831. Fig. 21.Basionym: Ascochyta heraclei Lib., Pl. crypt. Ard., Cent. 1: no. 51.1830.≡ Cyl<strong>in</strong>drosporium heraclei (Lib.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl. 115, I: 378. 1906 [non Oudem. 1873, nec Ellis & Everh. 1888].≡ Phloeospora heraclei (Lib.) Petr., Annls mycol. 17: 71. 1919 [non (Lib.)Maire, Bull. Soc. Mycol. France 46: 241. 1930].≡ Cyl<strong>in</strong>drosporium umbelliferarum Wehm., Mycologia 39: 475. 1947.nom. nov.= Sep<strong>to</strong>ria heraclei-palmati Maire, Bull. Soc. Mycol. France 21: 167. 1905.Description <strong>in</strong> planta: Symp<strong>to</strong>ms leaf spots numerous but small,irregular <strong>in</strong> outl<strong>in</strong>e, best visible on the upper side of the leaf,<strong>in</strong>itially yellowish or ochreous, later becom<strong>in</strong>g pale <strong>to</strong> dark brown,<strong>in</strong> places white due <strong>to</strong> loosen<strong>in</strong>g of the epidermis. Conidiomatapseudopycnidial, hypophyllous, one, rarely up <strong>to</strong> three <strong>in</strong> each260

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