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Reconsideration of Protocrea (Hypocreales, Hypocreaceae)

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Reconsideration of Protocrea (Hypocreales, Hypocreaceae) Iyeologio. 100(6), 2008, pp. 962-984. D( )I: 10.3852/08-101 2008 by The Nlv( ( logic1 Smivtv of America, I .awrcncc, KS 6604 1-8897 Reconsideration of Protocrea (Hypocreales, Hypocreaceae) \Valter M. Jaklitsch I tNtRt)tCtI()N 1icu/(y (:erilre for ,Syslematic Ru/any, ( Tn ive,/yrv of Vienna, Rennweg 74, A-1030 1 'len na. Austria Iivpocrea t ypically consists of a teleomorph, the sttonla of which is usuall y no more than a few Kadri POldmaa millimeters in diameter and includes priimtrily ?vatural 1-lislory Museum, Unit 'irsily u//ann, pseiidoptrenchytmitous tissue, bicellular ascospores Vanemnuise 46, E.E-51014, Tartu. /toiiia that disarticulate at the septum and a Tuieliodenna Gary j. Samuels Pers. anaiiiorph with green or, less frequently. 1/nited Stales Department a! Agriculture, .-tgricn/tura/ colorless (white in mass) con idia. however several Research Service, Sys/emalic Myco/u&çv awl Microbioluev described species vary troll) this stereotype in having Laboratory, Room 304, B-01 ]A, I7ARC-W, Beltsville, effused stroniata or a subiculum and/or having Maryland 20705 acremoniuln-. vetticilliuin- or gliocladium-like ana- rnorphs. Segregate genera have been proposed for some of these species, including Prolorrea and Abstract: The genus Protoerea is redefined, based on Arac/tnocrea Moravec, but in the absence of critical holotype and fresh specimens of its t P. ype species study, the paucity of' Specimens and the lack of farinosa, using morphology of tcleoinorph and cultures that elucidate ananiorphs while providing anamorph and phvlogenetic analyses of rph2 sequelic- molecular phvlogenetic infoimal ion correct applica- es. Data based on currentl y available specimens tion of these genetic names and assignment of species suggest the existence of three well defined and thiee has been haphazard. still unnamed Species Apart from the type, P. Petch (1937) erected the genus Proloerea including /arinosa, none of the species originally included are IIvpoerea Jarinosa Berk. & Broome, H. de/icatula Tul. Proloerea accepted in the genus. Species of are and H. slipata (Lib.) Fuckel. He did not indicate a characterized by perithecia formed in or on a type species. Moravec (1956) lectorvpified the genus sitbiculiun. bicellular ascospores that disarticulate at With P. fiirinosa, while retaining P. delicalnia and the septum while still in the ascus and by anamorphs removing P. st/pa/a to his new genus A ,achnoerea, the belonging to Ghoeladiu in sensu stile/u. For Hpocrea latter characterized b y disarticulating biconical asco- formosa sensu auci. the new species H. deci/nens is spores. Pôldmaa (2000) provided nioleculai phivloge- 1-Ipocrea pal/ida is introduced. recognized as a species netic evidence that the t y pe species of A rae/i noerea, A. Prooerea. of It is closely related to P. fti i/nasa, stipa/a, having a verticihlium-like ananiorph ( Pñl(niaa morphologicall Pro- y, phvlogeneticallvaiid by habit. 1999), is distinct hom flypoerea, including H. pal/ida. toerea i/h loensis is described here as the sister taxon Jlvpoerea (fr//ca/u/a in contrast belongs to J1ypoerea, Of P. fiirmnosa found in the USA. All species are based on gene sequences and a verticilhium-like polyporicolous, with the principal hosts Ske/etoculis (Trichoderina sect. Hvpocreanuin) anamorph (\V. nivea for P. farmnosa and P. illinoinsis, and species of Jaklitscli, unpubl) and will be treated elsewhere. Oligoporus/ iyromn 1ce.c pal/ida. for P. In addition to Overton oi al (2006h) epitvpiiied Protoerea, but as hosts the main differences among these species are a we will demonstrate this epitvpification was based on stronger (orange) pigmentation of perithecia and llliSidlell(ifiedl material and must be overturned. I. pal/ida suhiculum in and a violaceous KOH Another common species similar to P. Jitrinosa, reaction in P. pal/ida and P. i/h noeiisis. P. far/i/usa is producing perithecia in a suhictilum and it Glioc/adi- known only from Europe with certain iv and P. urn anamorph , and occurring 01) polypores is illinoènsis only from the USA, while P. pa//ida is I-Ipoerea pal/ida. Based on I,SLI ribosomal DNA probably cosmopolitan. Putative synonym y of some sequences, the species was found to fall outside similar species is discussed. Hypoerea in the Hvpocreaceae, bitt no generic Key wards: Ascomvcetes, Char/aim in. I-Iypocrra, redisposition has been suggested for it (Rehner and I-lvpocreales, ITS, LSU, morphology, phvlogcnv, Samuels 1994, lOklinaa ci al 1999, Pôldmaa 2000). )/)h2, sequence analysis, systematics, lefi The main objectives of this study are (i) the correct interpretation, conceptual definition and description of the genus Protociea and its type species P. fit rinosa, Accepted for pi,htic,itioti 29 July 2008. (ii) to determine the phvlogenetic l)OsitiOli of (.ii-poiuIiiig uohoi, I-I,1:re, \vIlt(I.J;Ik]INIhu( Univic,;u,:tI Protoerea and! Hypoerea /.iaiiida and (iii) to describe JANI tTS( :1-i El Al,: RECONS! ItERATION OF P/?OJoCJU.\ 963 tite lnngiis iiitetpieted as I1rp0re(1 faeino.ca b recent nucle;u tI)N,\, coittaining the I 151 and 2 legions. was authors as a new species of II/ocrea. amplified by PUP. with the piitiiet combinations SR(iR and I.R1 (\,lute el al 1990) oi with ITS4 (White ci al 1990) and ITS I F (Gaides and Bi-titis 1993). LSU rDNA w;ts amplified NL\lFIRiAI,S ANI) ME I bIts With the latter piiliter in couihmatiou xvith LRS (http:// ssww.hiologs .diike.edtt/ftuigt/mv( olab/prittiei-shitrn). A 1.SO/OICS and s/ieruncos.—I Sol ates and (enBiikoces.sioii 1.3 kb fiagtiient of the te/1 numbers for ITS. I Sf.. rp1)2 and IeJJ sequences in this study gene encoding tiatislatioti are listed (T \tti ,t, I ) . Isolates giver! as C. P.K. ( Kubicek) are elongation factor 1 alpha was amplified with the primer IOU! those niaiiit,ained ill (ollection of ilic Institute of EEl 728F and TEFI LLEi-ev Uaklitscl i et a] 2005. 2006). This Chemical Fngiiieenug. Vienna I. niveisitv of Iechnologv. fragtnent includes the f'ortrthi atid the filth introils and a those listed as (;.J.S. are those maintained at the I. SDA-ARS. part of the last large ex ui.A 0.9 kb fra gmentt of' RNA Svste tnatic Nis cologv and Microbiology I ala natorv, Belts- polvtnerase II stthtintt B ()1)1)2) was amplified with the ville, Macslat id midid those given as TF( are maintained at primer pair IRP112-5f mid I1UPB2-7et (Lin ci ;ih 1999). PUP. file f.Tttiveisjts of Life Sciences, Tartu, Estonia. Represen ta- pioclncis either ssete purified willi the QI,\quick Kit tOe isola es haxc beet i deposited at the ( eti t raalhoteau 5(0W (QI.\CEN) according to the manufacturers instrllclions Schinunelcrdtntes, Utrecht. The Netherlands (CBS). Spec- or with ;tti en/vulatic PUP. cleanup (WeiIe et al 199-I). For ittieus were deposited ill Heti)atmni of the Institute ol the latter 20 (tI. P( R reactions wei-e digested with 10 it Botany, Um yetsitv of Vienna. Austria (\VU), ill the U.S. exotiuclease I (Fetnientas.St I,eott-Roi, BRD) and 2 it calf National Fungus Collections, Beltsville, Maryland (RN), or intestine alkaline phosphi;tt;tse ( Feit tiet i tas) for IS min at the Estontan tiiiseisit y of Life Sciences, Tarot, Estonia 37 C. hi dlowed by ;tt i en /vinee cleact oat ion step at 85 C for 15 tutu . DNA was cycle-sequenced with the All PRISM Big Single-ascospoie isolates (( .P.K.., CBS strains) were Dye Teinunator C ycle Scr1ueiicitig Ready Reaction kit v. 3.1 pnp;tred 110111 I resh sj lecililCi 15 of / JYJJO(tea Strut! lata as (.\pplicd Biosyst (ms. \\atli i igton ) With the same ])I i mets as described hs Jaki i tscb i et a] (2006) or it mass of asci spores ill PUP., or with ITS and IIS5 (White et at 1990) for 115, or was isolated Olito 2% Ml,r\ (IFC strains). Cult ines wit Ii the rntertial printers 5-CC( ;lU,\(T/ )TT ( AICAA- designated G.J.S. sveie isolated vitli the use of it mictoina- G.\ACATG-3 and s-I"I-(x;C, GTC; ICCAICIT GITC-3 for ni pu I ;ttor oil thea] agar (Dilco ) snpl)lenie it ed with 2% fiji. ;u t d all automated DNA sequencercct- (,\Bl Genetic dextrose (C\ID). Analvzcrs Applied Biosvstcms) - (;!1nt/ (I1a?(1(/e?1zau/ofl.—Strtifl5 were cultivated oil i1o/ecn far Ju/ivfoginetic 0110lv,ve,s, —Seqiteuce ftagnieii ts were f(i nn in ea] agar, Sigma, St Louis,)uiS, Missouri) supplemented atialvzed and assembled ssithi SeqNlan Pro (lasergene. itli 2% (w"s) dextrose). PD,\ (potato-dextroseagar. l)N\STAR, Madisoti, Wisconsin) or Sequetichet 1.7 (G(,ne Merck, Darmstadt, Getniaiiv) and 2% ME,\ (malt extract Codes, Anti Arbor. Nlichigait). DNA sequences were agar. Merck) all ill led s atet. Growth took place at st d,tnitted to ( ;eti Batik. Alignmentsieti ts wet-c petiorined with 25 ( (alternating 12 h cool white fluorescent light and 12 It MAFFT v 6.240 ( Katoh ci al 2005), hd]ossed bs man tial dtikitess). adjustments with Genedoc 2.6 (Nicholas et ill 1997). Maxi ti utti paisitii! ins (NIP) atialvses sseie conducted in S Ioi/dinln i,"ual ob,sc,ivilia,i . of the—Structures ait;ti n (lii)] I prp 4.01)10 (Swolfot-d 2002) wiiii 110)01) Iietitistic were exami it ed , itteasured and photogiapl IC I on it corn- searches with randoni taxou addition sequences arid 11I pottt1d microscope horn ctiltuies glow!! Oil MD or Pl).\ at hr-itichi swapping. The cotihidetice Of bi;uicliing was assessed 25 C oil plates under low magnification and after with bootstrap resainpliig (bs) : 1000 replicates, each with mounting ill KOl I.
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