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

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Verkley et al.Fig. 32. Septoria petroselini. A, B. Colonies CBS 109521 (15 °C, nUV). A. On OA. B. On MEA. C, D. Conidia on OA (CBS 109521). E. Conidia and conidiogenous cells in planta(CBS H-21166). F. Conidia on OA (CBS 182.44). G. Conidia on OA (CBS 109521). Scale bars = 10 µm.Hosts: Petroselinum crispum (syn. Apium petroselinum), otherPetroselinum spp. and Coriandrum sativum (Priest 2006).Material examined: Netherlands, Prov. Utrecht, Baarn, garden Eemnesserweg 90,on living leaves of Petroselinum crispum, 29 Mar. 2001, H.A. van der Aa 12642,CBS H-21166, living culture CBS 109521; Laren, on living leaves of P. sativum, June1944, S. Dudok de Wit s.n., living culture CBS 182.44 = IMI 100279, dried specimenof culture on CMA, CBS H-18128.Notes: CBS 182.44, isolated from Petroselinum sativum, producesconidia 29–49 × 1–2 µm, and this range of sizes agrees withthose given for S. petroselini by most authors [26–45(–52) ×(1–)1.5–2 µm cf. Priest 2006; 16–46 × 1–2 mm cf. Jørstad 1965on Petroselinum]. In contrast, the conidia in the collection on P.crispum (CBS H-21166), as well as in the isolate CBS 109521derived from it, were up to 80 µm long and 2.5 µm wide, and thepycnidia were also larger than described for S. petroselini, for whichthis material was initially identified as S. apiicola, but the moleculardata provide evidence that it also belongs to S. petroselini. Thematerial is 100 % homologous on ITS, Act, RPB2 and EF, and99.7 % on Cal with CBS 182.44. The range of conidial sizes for S.petroselini is therefore expanded here, although it should be notedthat the conidia formed in vitro are not over 65 µm in length in thematerial available. The ITS sequence of S. anthrisci is distinct fromthat of S. apiicola, but identical to that of S. petroselini and otherspecies. Septoria anthrisci can be distinguished from S. petroseliniby the Act, EF and RPB2 sequences.Septoria phlogis Sacc. & Speg., in Sacc., Michelia 1: 184.1878 [as “phlocis”; non Ellis & Everh., in G. Martin, J. Mycol.3: 85. 1887; nec P. Syd., Mycoth. March., Cent. 18, no 1757;Cent. 23, no 2278. 1887; later homonyms]. Fig. 33.Description in planta: Symptoms leaf lesions developing in areasof the leaf lamina that first turn yellow, indefinite or delimited bydarkening veinlets, hologenous, pale to dark brown. Conidiomatapycnidial, epiphyllous, numerous, semi-immersed to immersed,subglobose to globose, dark brown to black, 100–160 µmdiam; ostiolum central, circular, initially 25–35 µm wide, laterbecoming more irregular and up to 70 µm wide, surrounding cellsconcolourous; conidiomatal wall 15–28 µm thick, composed of anouter layer of isodiametric to irregular cells mostly 5–9 µm diamwith pale brown cell walls up to 2 μm thick, and an inner layer ofhyphal to isodiametric cells 3–5 μm diam with thin, hyaline walls.Conidiogenous cells hyaline, discrete or integrated in 1–2-septateconidiophores up to 22 µm long, cylindrical, or narrowly to broadlyampulliform, holoblastic, often proliferating percurrently withindistinct annellations as well as sympodially, 5–7.5(–8) × 2.5–4(–5) µm. Conidia cylindrical, filiform, straight to slightly curved,278
A new approach to species delimitation in SeptoriaNotes: Priest (2006) described the conidia of S. phlogis as filiform,1–4-septate, straight to curved, (35–)50–73 × (1–)1.5–2 µm,hyaline, with a truncate base and obtuse apex. He accepted S.divaricatae as a separate species, with Phlox drummondi (syn. P.divaricata) as the only known host plant, and S. drummondi as asynonym. Septoria divaricatae has similarly shaped but smallerconidia than S. phlogis, 1–3-septate, (13–)25–40(–45) × 1–1.5µm. The overlap in length of the conidia of the two is minimal, atleast on the host plant, indicating that they might be truly separatetaxa. Several other authors have also accepted S. divaricatae asa distinct entity (Teterevnikova-Babayan 1987, Muthumary 1999).However, Jørstad (1965) considered S. divaricatae a synonym ofS. phlogis, and also S. phlogina. Both S. phlogis and S. divaricataeoccur on P. drummondi and this may have contributed to theconfusion. Investigations based on fresh material on different Phloxspecies, and studies of cultures derived thereof, as well as typematerial of the names mentioned above, will be required in order tosettle the complicated taxonomy of Septoria on Phlox.Molecular identification of S. phlogis is straight-forward, as allprotein-coding genes investigated here, particularly Btub, Cal andRPB2, show unique diagnostic sequences. Septoria epambrosiae(CBS 128629, 128636) is a sister species to S. phlogis. Septoriaepambrosiae is a pathogen of Ambrosia artemisiifolia (Asteraceae),which today is the prime cause of hay fever in many areas wherethis weed occurs.Fig. 33. Septoria phlogis. Conidia and conidiogenous cells in planta (CBS H-21198).Scale bars = 10 µm.narrowly rounded to somewhat pointed at the apex, attenuatedgradually or more abruptly towards the narrowly truncate base,(0–)1–3(–4)-septate, not constricted around the septa, hyaline,containing minute oil-droplets and granular material in the livingand rehydrated state, (22–)32–50(–60) × 1.5–2 µm (rehydrated;living, 2–2.5 µm wide). Sexual morph unknown.Description in vitro: Colonies on OA 15–18 mm diam in 19 d, with aneven, glabrous, buff to rosy-buff margin; colonies spreading, plane;immersed mycelium variably pigmented over sectors, usually eitherbrownish olivaceous, or cinnamon to saffron (honey with a reddishhaze); aerial mycelium scanty, white, locally forming a diffuse woollyfloccosemat; reverse olivaceous-black and cinnamon or saffron.Colonies on CMA 13–18 mm diam in 19 d, as on OA. Colonies onMEA 12–17 mm diam in 19 d, with an even, glabrous, buff margin;colonies spreading, the surface mostly plane, only somewhatelevated or folded towards the centre; immersed mycelium mostlydark salmon to olivaceous-black, covered by a dense, appressedmat of woolly, mostly white to faintly rosy-buff aerial mycelium; anochreous pigment diffuses into the surrounding medium; reversemostly sienna or blood colour, with an ochreous to saffron margin.Colonies on CHA 12–18 mm diam in 19 d, as on MEA.Hosts: Phlox spp.Material examined: Netherlands, Prov. Noord-Holland, Enkhuizen, on living leavesof Phlox sp., 6 Sep. 1949, J.A. von Arx s.n., CBS H-4862; Prov. Utrecht, Baarn,Cantonspark, on living leaves of Phlox sp., 27 Aug. 1999, G. Verkley 911, CBSH-21198, living culture CBS 102317; same substr., Jan. 1932, D. Moll s.n., livingculture CBS 312.32; Garden in Baarn, same substr., 16 Oct. 1990, H.A. van der Aa10919, CBS H-18130, living culture CBS 577.90; same substr., loc., 27 Aug. 1997,H.A. van der Aa 12302, CBS H-18131.Septoria polygonorum Desm., Annls Sci. Nat., sér. 2,Bot.17: 108. 1842. Fig. 34.≡ Spilosphaeria polygonorum (Desm.) Rabenh., Herb. Mycol. II, no.442a. 1856.Description in planta: Symptoms leaf spots small, circular,hologenous, ochreous to brown, sharply delimited by a dark redbrownzone. Conidiomata pycnidial, mainly epiphyllous, severalto many developed in each leaf spot after some time, subgloboseto lenticular, not protruding strongly, brown to almost black, 50–120 µm diam; ostiolum central, initially circular and 25–45 µmwide, surrounding cells concolorous to somewhat darker brown;conidiomatal wall about 10–25 µm thick, composed of angularcells 2.0–6.5 µm diam, the outermost cells pale yellowish brownwith somewhat thickened walls, the inner cells thin-walled.Conidiogenous cells hyaline, discrete, narrowly or broadlyampulliform with a relatively wide neck, holoblastic, often firstproliferating sympodially, and later also percurrently 1–several timeswith distinct annellations, 5–10(–14) × 3.–5.5(–6.5) µm. Conidiafiliform to filiform-cylindrical, straight or slightly curved, or flexuous,attenuated gradually to a narrowly rounded to pointed apex,attenuated more abruptly towards the truncate base, 1–4-septate,not or only inconspicuously constricted around the septa, hyaline,contents with several minute oil-droplets and granular material ineach cell in the living state, with inconspicuous oil-droplets andgranular contents in the rehydrated state, (17–)22–45(–53) × 1.5–2µm (living; rehydrated, 1.2–1.8 µm wide). Sexual morph unknown.Description in vitro: Colonies normally slow-growing, but sometimeswith fast-growing sectors (diam including these between brackets)on all media except MEA. On OA 3–5 [6–7] mm diam in 2 wk [6–7(22–30) mm in 6 wk], the margin regular, glabrous, colourless;colonies spreading, plane, immersed mycelium olivaceous-black,but grey-olivaceous to greenish grey in faster growing sectors thatsometimes develop from typically slow-growing colonies; aerialmycelium generally absent or very scanty, but woolly-floccosewww.studiesinmycology.org279
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Page 18 and 19: Verkley et al.CaryophyllaceaeRutace
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Page 38 and 39: Verkley et al.wk (15-18 mm in 7 wk)
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Page 42 and 43: Verkley et al.Fig. 16. Septoria cuc
Page 44 and 45: Verkley et al.Fig. 17. Septoria dig
Page 46 and 47: Verkley et al.Baarnsche bos, ex lea
Page 48 and 49: Verkley et al.Septoria galeopsidis
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Page 52 and 53: Verkley et al.Fig. 23. Septoria lac
Page 54 and 55: Verkley et al.much more profound di
Page 56 and 57: Verkley et al.Fig. 26. Septoria lyc
Page 58 and 59: Verkley et al.Fig. 27. Septoria lys
Page 60 and 61: Verkley et al.to very dark dull gre
Page 62 and 63: Verkley et al.Fig. 30. Septoria nap
Page 64 and 65: Verkley et al.Fig. 31. Septoria par
Page 68 and 69: Verkley et al.Fig. 34. Septoria pol
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Page 74 and 75: Verkley et al.to 60 µm diam, lacki
Page 76 and 77: Verkley et al.Fig. 38. Septoria sis
Page 78 and 79: Verkley et al.on OA, but olivaceous
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Page 82 and 83: Verkley et al.ampulliform to almost
Page 84 and 85: Verkley et al.diam; Conidiogenous c
Page 86: Verkley et al.Notes: The type mater
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