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Black-foot disease of grapevine: an update on taxonomy, epidemiology and management strategies

Agustí Brisach, Carlos,Armengol Fortí, Josep

Abstract

[EN] Black-foot is one of the most destructive grapevine trunk diseases in nurseries and young vineyards, causing necrotic root lesions, wood necrosis of the rootstock base, and a gradual decline and death of grapevines. Causal agents of the disease are included into the genera Campylocarpon, "Cylindrocarpon", Cylindrocladiella and Ilyonectria. Recent taxonomical studies of Neonectria and related genera with "Cylindrocarpon"-like anamorphs based on morphological and phylogenetic studies, divided Neonectria into five genera. Thus, the current taxonomical position and classification of the causal agents of black-foot disease, mainly "Cylindrocarpon"/Ilyonectria, comprises one of the main topics of this review. The review also provides an update on geographical distribution, epidemiology and management strategies of the disease.

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245 www. up ess.com/pm ISSN (p in ): 0031-9465 © Fi enze Uni e si y P ess ISSN (online): 1593-2095 Phy opa hologia Medi e anea (2013) 52, 2, 245−261 Co esponding au ho : J. A mengol Fax: +34 963879269 E-mail: [email p o ec ed].es REVIEW Black- oo disease o g ape ine: an upda e on axonomy, epidemiology and managemen s a egies Ca los aGUsTÍ-B IsaCH and Josep a MeNGol Ins i u o Ag o o es al Medi e áneo, Uni e sidad Poli écnica de Valencia, Camino de Ve a s/n, 46022-Valencia, Spain Summa y. Black- oo is one o he mos des uc i e g ape ine unk diseases in nu se ies and young ineya ds, causing nec o ic oo lesions, wood nec osis o he oo s ock base, and a g adual decline and dea h o g ape ines. Causal agen s o he disease a e included in o he gene a Campyloca pon, “Cylind oca pon”, Cylind ocladiella and Ily- onec ia. Recen axonomical s udies o Neonec ia and ela ed gene a wi h “Cylind oca pon”-like anamo phs based on mo phological and phylogene ic s udies, di ided Neonec ia in o i e gene a. Thus, he cu en axonomical po- si ion and classi ica ion o he causal agen s o black- oo disease, mainly “Cylind oca pon”/Ilyonec ia, comp ises one o he main opics o his e iew. The e iew also p o ides an upda e on geog aphical dis ibu ion, epidemiol- ogy and managemen s a egies o he disease. Key wo ds: Campyloca pon, “Cylind oca pon”, Cylind ocladiella, Ilyonec ia, Vi is ini e a. In oduc ion Black- oo disease o g ape ines is a se ious dis- ease in mos wine and g ape-p oducing egions o he wo ld, pa icula ly in nu se ies and young ineya ds (Halleen e al., 2006a). The causal agen s a e included in o he gene a Campyloca pon, “Cylin- d oca pon”, Cylind ocladiella and Ilyonec ia (C ous e al., 1993; Halleen e al. 2004; Halleen e al., 2006b; Sch oe s e al., 2008; Cha e i e al., 2011; Cab al e al., 2012a, c; Lomba d e al., 2012). This disease was i s desc ibed in 1961 (G asso and Magnano Di San Lio, 1975), and o e he las decade, i s incidence has inc eased signi ican ly in mos g ape ine p oduc- ion a eas o he wo ld (Halleen e al., 2006a; Alaniz e al., 2007). Al hough hese pa hogens usually mani es on ma u e g ape ines, hey ha e also been equen ly isola ed om symp oma ic o asymp oma ic oo - s ock mo he -plan s, oo ed oo s ock cu ings, bench-g a and young g a ed ines in di e en g ape ine p oduc ion a eas a ound he wo ld, be- ing conside ed he mos common pa hogenic ungi associa ed wi h young nu se y ines (Rumbos and Rumbou, 2001; Halleen e al., 2003; Fou ie and Hal- leen, 2004; Oli ei a e al., 2004; A oca e al., 2006; Dub o sky and Fab i ius, 2007; Halleen e al., 2007). Mo eo e , i is well known ha hese pa hogens a e common in he soil causing in ec ion o g a ed ines a e some mon hs o g ow h in nu se y soils (Hal- leen e al. 2003, 2007; Cha e i e al., 2011). Cha ac e is ic symp oms o black- oo disease in- clude a educ ion in oo biomass and oo hai s wi h sunken and nec o ic oo lesions (Rego e al., 2000; Halleen e al., 2006a; Alaniz e al., 2007, 2009; Ab eo e al., 2010). In some cases he oo s ock diame e o old- e ines is hinne below he second ie . To compen- sa e o he loss o unc ional oo s, a second c own o ho izon ally g owing oo s is some imes o med close o he soil su ace. Remo al o oo s ock ba k e eals black discolo a ion and nec osis o wood is- sue which de elops om he base o he oo s ock (Figu es 1A, 1B). The pi h is also compac ed and dis- Phy opa hologia Medi e anea 246 C. Agus í-B isach and J. A mengol colo ed (Scheck e al., 1998b; La ignon, 1999; Fou ie and Halleen, 2001; Halleen e al., 2006a). Ex e nal symp oms show educed igou wi h small-sized unks, sho ened in e nodes, une en wood ma u i y, spa se oliage, and small lea es wi h in e einal chlo osis and nec osis (Figu es 1C, 1D). Field symp oms o black- oo disease a ec ed ines a e equen ly indis inguishable om hose o caused by Pe i disease (Scheck e al., 1998b; Rego e al., 2000; Halleen e al., 2006a; Alaniz e al., 2007, 2009; Ab eo e al., 2010). When young ines a e in ec ed, dea h oc- cu s quickly, ne e heless as he ine ages, in ec ion esul s in a mo e g adual decline and dea h migh only occu a e a yea (Guble e al., 2004). Disease symp oms on ma u e ines (5 yea s and olde ) a e no iced ea ly in he g owing season. A ec ed ines achie e poo new g ow h, ail o o m shoo s a e win e do mancy, and die by mid-summe . O en shoo s also d y and die du ing he summe . Vines wi h educed ege a i e g ow h also die du ing he subsequen do man win e pe iod (Halleen e al., 2006a). Causal agen s Taxonomy and dis ibu ion The common name black- oo disease was p o- posed by Scheck e al. (1998b), o designa e he dis- ease caused by “Cylind oca pon” des uc ans (Zinnsm.)          B C D A Figu e 1. A, black discolo a ion and nec osis o wood issue which de elops om he base o he oo s ock, cha ac e is ic o black- oo disease; B, longi udinal sec ion o a oo s ock showing da k-b own o black discolo a ion; C, Un-sp ou ed g ape ine p opaga ion ma e ial in a g ape ine nu se y; D, g ape ine plan s showing s unded g ow h, educed igou and e a ded sp ou ing in a young plan a ion. 247 Vol. 52, No. 2, Augus , 2013 An upda e o black- oo disease o g ape ine Schol en and “C.” ob usispo um (Cooke & Ha kn.) Wollenw., which we e he wo species adi ionally epo ed as he causal agen s o basal o o oo ne- c osis on g ape ines. Ne e heless, his disease was al eady named as “pied noi ” in F ench language since 1969, because o he p esence o black nec osis on he base o diseased oo s ocks (Badou , 1969). The i s epo o “C.” des uc ans on g ape ine was made in F ance in 1961 (Malu a and La ignon, 1991). Since hen, i has been isola ed om diseased ines in I aly (G asso, 1984), Po ugal (Rego, 1994), Cali o nia (Scheck e al., 1998b), A gen ina (Ga ica e al., 2001), Ge many (Fische and Kassemeye , 2003), Pennsyl ania (Gugino and T a is, 2003), New Zea- land and Sou h A ica (Halleen e al., 2004), B azil (Ga ido e al., 2004) and Canada (Pe i e al., 2011). “Cylind oca pon” ob usispo um, has also been epo - ed o p oduce black- oo symp oms on g ape ine in Sicily (G asso and Magnano di San Lio, 1975) and Cali o nia (Scheck e al., 1998a). The gene ic name “Cylind oca pon” was in o- duced in 1913 by Wollenwebe o anamo phs be- longing o Nec ia sec ion Willkommio es Wollenw. This sec ion included species wi hou chlamydo- spo es. Few yea s la e , in 1917, Wollenwebe ex- panded he concep o “Cylind oca pon” o include species o ming mycelial chlamydospo es in cul u e, being “C.” des uc ans he mos impo an membe o his g oup (B ay o d, 1993). In 1966, Boo h spli he genus in o ou g oups based on he p esence o absence o mic oconidia and chlamydospo es: (i) “Cylind oca pon” magnusianum (Sacc.) Wollenw., which was he anamo ph o he ype species o Neo- nec ia, (ii) “C.” cylind oides Wollenw., which was he ype species o he genus “Cylind oca pon”, (iii) “C.” des uc ans, which was he anamo ph o Neonec ia adicicola, and (i ) membe s o “Cylind oca pon” spe- cies p edominan ly connec ed wi h eleomo phs o he ‘Nec ia’ mammoidea g oup (B ay o d, 1993; Hal- leen e al., 2006a). “Cylind oca pon” ob usispo um was o iginally desc ibed om he USA (Cali o nia) as oc- cu ing on Acacia sp., whe e i was obse ed o o m mac oconidia and chlamydospo es (Boo h, 1966). “Cylind oca pon” ob usispo um s ains iden i ied by Boo h (1966) o igina ed om a b oad ange o hos plan s in Eu ope, New Zealand, No h Ame ica, and, a leas pa ly, o med mic oconidia. T adi ionally, ep esen a i es o all ‘Nec ia’ g oups wi h “Cylind oca pon” anamo phs we e ans e ed in o Neonec ia (Rossman e al., 1999; Man i i e al., 2001; B ay o d e al., 2004). Man i i e al. (2001) and B ay o d e al. (2004) analyzed mi ochon- d ial small subuni (SSU) ibosomal DNA ( DNA) sequence da a o some o he species and concluded ha he Neonec ia/“Cylind oca pon” species g ouped oge he by his eclassi ica ion we e monophyle ic. Howe e , hese au ho s also ound ha his o e all Neonec ia/“Cylind oca pon” clade included dis inc subclades co esponding o a leas h ee o he ou g oups delinea ed by Boo h (1966). Signi ican mo- lecula a ia ion among axa wi h “Cylind oca pon”- like anamo phs was ound by Sei e e al. (2003) in a s udy on ungi causing oo o o ginseng (Panax quinque olius L.) and o he hos s. The dend og ams in his s udy, based on pa ial β- ubulin gene (TUB), and nuclea ibosomal in e nal ansc ibed space (ITS) egion sequences, sugges ed ha subclades including (i) Neon. adicicola, which consis ed o nume ous phylogene ically dis inc uni s, (ii) Neon. mac oconidialis (Samuels & B ay o d) Sei e , and (iii) a subclade comp ising wo dis inc isola es, one om V. ini e a in On a io, Canada and he o he om Pi- cea sp. in Quebec, Canada, we e monophyle ic. O h- e “Cylind oca pon” species appea ed o be excluded om his monophyle ic g oup (Halleen e al., 2006a). Signi ican a ia ion in cul u al and mo phologi- cal cha ac e s was obse ed among “Cylind oca pon” s ain isola es om g ape ines in nu se ies and ine- ya ds o Sou h A ica, New Zealand, Aus alia and F ance, which we e mo phologically and phyloge- ne ically cha ac e ized by Halleen e al., (2004). Thus, hese au ho s desc ibed a no el species, “C.” mac o- didymum Sch oe s, Halleen & C ous, also associa ed wi h black- oo disease o g ape ines. Since hen, his species has been epo ed in Cali o nia (Pe i and Guble , 2005), Po ugal (Rego e al., 2005), Chile (Auge e al., 2007), Spain (Alaniz e al., 2007), U u- guay (Ab eo e al., 2010), no heas e n Uni ed S a es and sou heas e n Canada (Pe i e al., 2011) and Tu - key (Özben e al., 2012). “Cylind oca pon” ob usispo um and “C.” mac o- didymum had been conside ed as wo di e en spe- cies associa ed wi h black- oo disease o g ape ines. Ne e heless, Halleen e al. (2004) sugges ed he pos- sibili y ha G asso and Magnano di San Lio (1975) and Scheck e al. (1998a) misiden i ied “C.” ob usispo- um and ha i was in ac “C.” mac odidymum. In his sense, Halleen e al. (2004) indica ed ha mac- oconidia o “C.” mac odidymum measu e [(26–)34– 36–38(–45)×(4–)5.5–6–6.5(–8) μm], whe eas hose o Phy opa hologia Medi e anea 248 C. Agus í-B isach and J. A mengol he ype o “C.” ob usispo um measu e (30–35×4–5 μm) (Cooke, 1884). Howe e , he shape o he mac- oconidia dis inguishes “C.” mac odidymum om he ype o “C.” ob usispo um, which Cooke (1884) de- sc ibed as ha ing conidia wi h ob use ends. Boo h (1966) desc ibed mac oconidia o simila shape in “C.” ob usispo um. Acco ding o Boo h, howe e , 2–3-sep a e mac oconidia o “C.” ob usispo um meas- u e (34–50×6–7.5 μm). “Cylind oca pon” ob usispo um isola es ob ained om Cali o nia o med pe i hecia when c oss-inocula ed wi h “C.” mac odidymum, gi ing u he e idence o suppo he misiden i i- ca ion heo y. This was also con i med by sequence compa isons (Halleen e al., 2006a). In 2005, Pe i and Guble con i med he p esence o “C.” mac odidymum in he USA, and concluded ha black- oo disease in Cali o nia is caused by “C.” mac odidymum and “C.” des uc ans (Pe i and Guble , 2005). Mo eo e , Halleen e al., (2004) es ablished a new genus, Campyloca pon Halleen, Sch oe s & C ous, which is “Cylind oca pon”-like in mo - phology, associa ed wi h black- oo disease o g ape ines. Species o his genus and membe s o he o me “Nec ia” mammoidea g oup, a e ex- cluded om Neonec ia/“Cylind oca pon”, because phylogene ic analyses e ealed ha hese spe- cies a e phylogene ically no closely ela ed o Neonec ia/“Cylind oca pon” gene a (Halleen e al., 2004; Sch oe s e al., 2008). F om his genus, wo spe- cies we e included as he causal agen s o black- oo disease: Campyloca pon ascicula e Sch oe s, Halleen & C ous, which has been epo ed in Sou h A ica (Halleen e al., 2004), B azil (Co eia e al., 2012), and Spain (Alaniz e al., 2011b) and Campyl. pseudo ascicu- la e Halleen, Sch oe s & C ous, which has been e- po ed in Sou h A ica (Halleen e al., 2004), U uguay (Ab eo e al., 2010), B azil (Co eia e al., 2012) and Pe ú (Ál a ez e al., 2012). As highligh ed be o e, “C.” des uc ans was o igi- nally iden i ied as he causal agen o black- oo dis- ease (Malu a and La ignon, 1991), bu he s a us o “C.” des uc ans as he causal agen o he disease was since hen ques ioned. In ac , Halleen e al. (2006b), compa ed “C.” des uc ans s ains isola ed om diseased g ape ines in F ance, New Zealand, Po ugal and Sou h A ica wi h “C.” des uc ans-like anamo phs ob ained om a ious he baceous o woody hos s. DNA analyses o hei ITS and TUB showed ha hese isola es we e gene ically iden i- cal wi h “C.” li iodend i J.D. MacDon. & E.E. Bu le , which was i s associa ed wi h oo o o ulip pop- la (Li iodend on ulipi e a L.) in Cali o nia by Mac- Donald and Bu le (1981). Thus, because hese spe- cies had iden ical sequences, “C.” des uc ans isola es collec ed om asymp oma ic o diseased g ape ines a ec ed by black- oo disease we e enamed as “C.” li iodend i, associa ing “C.” des uc ans only wi h oo o on o he he baceous o woody hos s (Hal- leen e al., 2006b). In addi ion, in o de o cla i y he axonomy o “C.” des uc ans causing black- oo in Cali o nia, Pe i and Guble (2007) also compa ed “C.” des uc ans isola es ob ained om g ape ines in Cali o nia wi h “C.” li iodend i isola es om Sou h A ica. All o hem we e iden ical, and consequen ly “C.” des uc ans isola es we e also enamed as “C.” li iodend i. This species has been la e epo ed as a black- oo pa hogen o g ape ine in Aus alia (Whi elaw-Wecke e al., 2007), Spain (Alaniz e al., 2007), B azil (Russi e al., 2010), I an (Mohammadi e al., 2009), Swi ze land (Casie i e al., 2009), U uguay (Ab eo e al., 2010) and no heas e n Uni ed S a es and sou heas e n Canada (Pe i e al., 2011). The ele- omo phs o “C.” li iodend i and “C.” mac odidymum we e desc ibed as Neonec ia li iodend i Halleen, Rego & C ous and N. mac odidyma Halleen, Sch oe s & C ous (Halleen e al., 2004, 2006b). In 2008, a new species associa ed wi h black- oo disease o g ape ines, “C.” paucisep a um Sch oe s & C ous, was desc ibed in New Zealand and Slo enia (Sch oe s e al., 2008). To da e, his species has been isola ed om a ec ed g ape ines in U uguay (Ab eo e al., 2010), Canada (O’Go man and Haag, 2011), Spain (Ma in e al., 2011) and Po ugal (Cab al e al., 2012a). Phylogene ic s udies ca ied ou in New Zea- land and Slo enia by Sch oe s e al. (2008), indica ed ha “C.” paucisep a um is he closes phylogene ic sis e - axon o “C.” mac odidymum and bo h species a e closely ela ed o he “C.” des uc ans-complex, which also includes “C.” li iodend i. Thus, a his momen , “C.” des uc ans, “C.” li i- odend i, “C.” mac odidymum, “C.” ob usispo um, “C.” paucisep a um, Campyl. ascicula e and Campyl. pseu- do ascicula e we e conside ed as he main species associa ed wi h young ines showing symp oms o black- oo disease in mos o g ape ine p oducing a eas wo ldwide. In addi ion, o he “Cylind oca pon” species ha e been associa ed occasionally wi h black- oo disease o g ape ine: “Cylind oca pon” didymum (Ha ing) Wollenw. in Canada (Pe i e al., 2011), “C.” olidum (Wollenw.) Wollenw. in Spain (De F ancisco 249 Vol. 52, No. 2, Augus , 2013 An upda e o black- oo disease o g ape ine e al., 2009) and “C.” olidum a . c assum Ge lach in U uguay (Ab eo e al., 2010). Cha e i e al. (2011) pe o med a phylogene ic s udy o Neonec ia, “Cylind oca pon” and ela ed gene a wi h “Cylind oca pon”-like anamo phs. Mo - phological and molecula phylogene ic analyses da a accumula ed o e se e al yea s ha e indica ed ha Neonec ia sensu s ic o and “Cylind oca pon” sensu s ic o a e phylogene ically congene ic, while Neonec ia sensu la o and “Cylind oca pon” sensu la o do no o m a monophyle ic g oup, sugges ing ha Neonec ia/“Cylind oca pon” ep esen s mo e han one genus. Thus, based on esul s o he phylogene - ic s udy, hese au ho s di ided Neonec ia in o i e gene a based on a combina ion o cha ac e s linked o pe i hecial ana omy and conidial sep a ion: Neonec ia/“Cylind oca pon” sensu s ic o (Boo h’s g oups 1 and 4), Rugonec ia, Thelonec ia (g oup 2), Ilyonec ia (g oup 3) and anamo ph genus Campy- loca pon. Acco ding o his, only Neonec ia has “Cylind oca pon” anamo phs, while he emaining gene a ha e “Cylind oca pon”-like anamo phs, and since hen a e e e ed o as “Cylind oca pon”. Con- sequen ly, “C.” li iodend i and “C.” mac odidymum we e included in o Ilyonec ia genus, wi h I. adici- cola as he ype species, and e-iden i ied as Ilyonec- ia li iodend i (Halleen, Rego & C ous) Cha e i & Salgado and I. mac odidyma (Halleen, Sch oe s & C ous) P. Cha e i & Salgado, espec i ely (Cha e - i e al., 2011). Mo eo e , Cab al e al. (2012a) we e able o deline- a e 12 new axa in he I. adicicola-complex, p e ious- ly known as he “C.” des uc ans-complex, by using a mul i-gene DNA analysis suppo ed by mo pho- logical cha ac e s. O he Ilyonec ia species wi hin I. adicicola-complex ha e been also ound associa ed wi h black- oo disease o g ape ine: Ilyonec ia eu o- paea A. Cab al, Rego & C ous, I. lusi anica A. Cab al, Rego & C ous, I. pseudodes uc ans A. Cab al, Rego & C ous and I. obus a (A.A. Hildeb .) A. Cab al, Rego & C ous, epo ed in Po ugal (Cab al e al., 2012a, 2012c). Ano he Ilyonec ia spp., I. i is has also been desc ibed in Po ugal (Cab al e al., 2012a), and iso- la es belonging o Neonec ia mammoidea g oup ha e also been associa ed wi h he disease in Canada (Pe - i e al., 2011). Soon he ea e , ollowing his s udy, Cab al e al. (2012c), demons a ed he exis ence o polymo phism in o I. mac odidyma-complex. This hypo hesis was in ag eemen wi h he esul s ob- ained by Alaniz e al. (2009), who al eady de ec ed ele an gene ic di e si y in “C.” mac odidymum by using in e -simple sequence epea (ISSR) echnique. Howe e , p e ious phylogene ic analysis showed low a ia ion in he la ge subuni (LSU) ibosomal DNA ( DNA), TUB and ITS sequences o “C.” mac o- didymum isola es ob ained om g ape ine in di e - en coun ies (Halleen e al, 2004; Pe i and Guble , 2005; Alaniz e al., 2007). Thus, in o de o cla i y his hypo hesis, Cab al e al. (2012c) pe o med a phylogene ic s udy o I. mac odidyma-complex by using ITS, TUB, his one H3 gene (HIS) and ansla- ion elonga ion ac o 1-α (TEF) sequence analysis. Consequen ly, six new species o Ilyonec ia (I. al- cace ensis A. Cab al, Oli ei a & C ous, I. es emocen- sis A. Cab al, Nascimen o & C ous, I. no ozelandica A. Cab al & C ous, I. o esensis A. Cab al, Rego & C ous, and Ilyonec ia sp. 1, I. sp. 2,) and I. mac o- didyma, which a e mo phologically a he simila , we e ecognised in o he I. mac odidyma-complex. All hese species ha e been epo ed in Po ugal, wi h he excep ion o I. no ozelandica which has been e- po ed in Sou h A ica, USA, New Zealand (Cab al e al., 2012a, 2012c). Recen ly, I. alcace ensis, I. mac o- didyma, I. no ozelandica, and I. o esensis ha e also been ound on g ape ines in Spain (Agus í-B isach e al., 2013a, 2013b). Rega ding, “C.” paucisep a um, i is no clea in which gene a i has o be included, al hough i is e y simila in mo phology o I. an hu iicola A. Cab al & C ous (Cab al e al., 2012a). Finally, ano he genus, Cylind ocladiella Boesew., which is also Cylind oca pon-like in mo phology, has ecen ly been associa ed wi h black- oo disease o g ape ines (Van Colle e al., 2005; Agus í-B isach e al., 2012; Jones e al., 2012). This genus was es ab- lished by Boesewinkel (1982) o accommoda e i e Cylind ocladium-like species p oducing small and cy- lind ical conidia. This decision was based on he ac ha species o Cylind ocladiella had di e en conidi- opho es b anching pa e ns, conidial shapes, dimen- sions, cul u al cha ac e is ics and eleomo phs om hose o Cylind ocladium (Van Colle e al., 2005; Lom- ba d e al. 2012). Since hen, se e al axonomic s ud- ies o hese ungi ha e elied on mo phologically and o lesse ex en on DNA sequence compa isons o he ITS and TUB gene egions, ecognizing nine species o Cylind ocladiella (C ous and Wing ield, 1993; Vic- o e al., 1998; Van Colle e al., 2005). Lomba d e al. (2012), ha e jus desc ibed 18 new Cylind ocladiella species based on mo phological and phylogene ic Phy opa hologia Medi e anea 250 C. Agus í-B isach and J. A mengol s udies employing ITS, TUB, HIS and TEF gene e- gions. Ne e heless, only wo species in o his genus ha e been ound associa ed wi h black- oo disease on g ape ines: Cylind ocladiella pa a (P.J. Ande son) Boesew., which has been epo ed in Sou h A ica (Van Colle e al., 2005), New Zealand (Jones e al., 2012) and Spain (Agus í-B isach e al., 2012) and Cyl. pe u iana (Ba ., J.L. Beze a & M.P. He e a) Boesew., which has been epo ed in Sou h A ica (Van Colle e al., 2005), Pe ú (Ál a ez e al., 2012) and Spain (Agus í-B isach e al., 2012). A lis o all “Cylind oca pon”/Ilyonec ia, Campylo- ca pon and Cylind ocladiella species, which ha e been epo ed associa ed wi h black- oo disease o g ape- ine and hei geog aphical dis ibu ion, is p esen - ed in Table 1. Mo phological and cul u al cha ac e iza ion “Cylind oca pon”/Ilyonec ia, Campyloca pon and Cylind ocladiella species ha e cha ac e is ic dis inc- i e mo phological and cul u al pa e ns (Figu es 2 and 3; Table 2). The anamo phs o “Cylind oca pon”/ Ilyonec ia p oduce abundan mic oconidia and chla- mydospo es. Mac o- and mic oconidia appa en ly a e p oduced om he same conidiopho es which a e 40–160 μm long, gene ally simple, unb anched o spa sely b anched, i egula ly o e icilla ely b anched, a ely densely b anched, and wi h cy- lind ical phialides. Mac oconidia a e s aigh o cu ed, hyaline, 1–3-sep a e, a ely > 3-sep a e [25– 50(–55)×5–7.5 μm], gene ally wi h a p ominen ba- sal o la e al abscission sca o hilum. Mic oconidia a e ellipsoidal o o oid, hyaline, 0–1-sep a e, wi h a la e al o basal hilum [3–15×2.5–5(–6) μm] (Figu es 2A, 2B). Chlamydospo es a e abundan , gene ally in e cala y, globose, single o in chains, becoming b ownish. In addi ion, colony mo phology on PDA is e y he e ogeneous (Figu e 2C). Ae ial mycelium is loccose o el ed, and he colou a ies om whi e o yellow o ligh o da k b own. The ma gin o he colony can be en i e, sligh ly lobula ed, o lobula ed (Figu es 3A–3H) (Boo h, 1966; Samuels and B ay- o d, 1990; Cha e i e al., 2011). Table 1. Fungal species which ha e been epo ed associa ed wi h black- oo disease o g ape ines and hei geog aphical dis ibu ion. Species Dis ibu ion Campyloca pon ascicula e Sch oe s, Halleen & C ous Sou h A ica (Halleen e al., 2004), Spain (Alaniz e al., 2011b) and B azil (Co eia e al., 2012). Campyloca pon pseudo ascicula e Halleen, Sch oe s & C ous Sou h A ica (Halleen e al., 2004), U uguay (Ab eo e al., 2010), B azil (Co eia e al., 2012) and Pe ú (Ál a ez e al., 2012) “Cylind oca pon” des uc ans (Zinssm.) Schol en F ance (Malu a and La ignon, 1991), I aly (G asso, 1984), A gen ina (Ga ica e al., 2001), Ge many (Fische and Kassemeye , 2003), Pennsyl ania (Gugino and T a is, 2003), B azil (Ga ido e al., 2004) and Canada (Pe i e al., 2011) “Cylind oca pon” didymum (Ha ing) Wollenw. Canada (Pe i e al., 2011) “Cylind oca pon” ob usispo um (Cooke & Ha kn.) Wollenw. Sicily (G asso and Magnano di San Lio, 1975) and Cali o nia (Scheck e al., 1998a) “Cylind oca pon” olidum (Wollenw.) Wollenw. Spain (De F ancisco e al., 2009) “Cylind oca pon” olidum a . c assum Ge lach U uguay (Ab eo e al., 2010) “Cylind oca pon” paucisep a um Sch oe s & C ous New Zealand and Slo enia (Sch oe s e al., 2008), U uguay (Ab eo e al., 2010) Canada (O’Go man and Haag, 2011), Spain (Ma in e al., 2011) and Po ugal (Cab al e al., 2012a) Cylind ocladiella pa a (P.J. Ande son) Boesew. Sou h A ica (Van Colle e al., 2005), New Zealand (Jones e al., 2012) and Spain (Agus í-B isach e al., 2012) (Con inued) 251 Vol. 52, No. 2, Augus , 2013 An upda e o black- oo disease o g ape ine Campyloca pon is simila o “Cylind oca pon”/Ily- onec ia, al hough Campyloca pon spp. p oduce mac- oconidia mos ly cu ed, while mic oconidia a e absen and chlamydospo es a e a e o also absen . Conidiopho es appea a ising la e ally om single o ascicula e ae ial hyphae o om c eeping sub- s a e hyphae, singly o in loose o dense agg ega es (Figu e 2D). Conidial heads o m pionno es-like ag- g ega es. Conidiopho e show a s ipe base o 16 μm wide, which bea se e al phialides o a penicillus o i egula b anches wi h e minal b anches bea ing 1 o se e al phialides. Mac oconidia a e as in Ilyonec- ia, bu ypically cu ed, and wi h up o 6 sep a, [(24–)35–60(–62)×6.5–9 μm], apical cell ob use, basal cell ob use o wi h inconspicuous hilum (Figu e 2E). Rega ding colony mo phology on PDA, ae ial myce- lium is abundan , co e ing he whole o sec o s o he colony, whi e o o -whi e o sligh ly b ownish, hick- ly co ony o el y, in e mingled wi h o gi ing ise o e ec whi e o b own hyphal s ands. This s ands some imes a e pa ly co e ed by o -whi e slime (Fig- u es 3I, 3J) (Halleen e al., 2004; Cha e i e al., 2011). Cylind ocladiella species p oduce hyaline, single, sub e icilla e, as well as penicillia e conidiopho es, Species Dis ibu ion Cylind ocladiella pe u iana (Ba ., J.L. Beze a & M.P. He e a) Boesew. Sou h A ica (Van Colle e al., 2005), Spain (Agus í-B isach e al., 2012) and Pe ú (Ál a ez e al., 2012) Ilyonec ia alcace ensis A. Cab al, Oli ei a & C ous Po ugal (Cab al e al., 2012c) and Spain (Agus í-B isach e al., 2013b) Ilyonec ia es emocensis A. Cab al & C ous Po ugal (Cab al e al., 2012c) Ilyonec ia eu opaea A. Cab al, Rego & C ous Po ugal (Cab al e al., 2012a) Ilyonec ia li iodend i (Halleen, Rego & C ous) Cha e i & Salgado F ance, New Zealand, Po ugal and Sou h A ica (Halleen e al., 2006b), Aus alia (Whi elaw-Wecke e al., 2007), Cali o nia (Pe i and Guble , 2007), Spain (Alaniz e al., 2007), I an (Mohammadi e al., 2009), Swi ze land (Casie i e al., 2009), B azil (Russi e al., 2010), U uguay (Ab eo e al., 2010), no heas e n Uni ed S a es and sou heas e n Canada (Pe i e al., 2011), Ilyonec ia lusi anica A. Cab al, Rego & C ous Po ugal (Cab al e al., 2012a) Ilyonec ia mac odidyma (Halleen, Sch oe s & C ous) P. Cha e i & C. Salgado Aus alia, F ance, New Zealand and Sou h A ica (Halleen e al., 2004), Cali o nia (Pe i and Guble , 2005), Chile (Auge e al., 2007), U uguay (Ab eo e al., 2010), no heas e n Uni ed S a es and sou heas e n Canada (Pe i e al., 2011), Po ugal (Cab al e al., 2012c), Tu key (Özben e al., 2012) and Spain (Agus í-B isach e al., 2013b). Ilyonec ia no ozelandica A. Cab al, Nascimen o & C ous Sou h A ica, USA and New Zealand (Cab al e al., 2012c) and Spain (Agus í-B isach e al., 2013a, b) Ilyonec ia pseudodes uc ans A. Cab al, Rego & C ous Po ugal (Cab al e al., 2012a) Ilyonec ia obus a (A.A. Hildeb .) A. Cab al, Rego & C ous Po ugal (Cab al e al., 2012a) Ilyonec ia o esensis A. Cab al, Rego & C ous Aus alia, Canada, New Zealand, Po ugal, Sou h A ica, Spain and USA (Cab al e al., 2012c; Agus í-B isach e al., 2013a, b) Ilyonec ia i is A. Cab al, Rego & C ous Po ugal (Cab al e al., 2012a) Ilyonec ia sp. 2 (Cab al e al., 2012c) Po ugal (Cab al e al., 2012c) Isola es belonging o Neonec ia mammoidea g oup Canada (Pe i e al., 2011) Table 1. Con inues. Phy opa hologia Medi e anea 252 C. Agus í-B isach and J. A mengol wi h p ima y and seconda y b anches. The phial- ides a e e minal, hyaline, in who ls o 2–4, wi h o wi hou ob ious colla e s. In gene al, s ipe is cen- ally a anged on conidiopho es, wi h a single basal sep um, e mina ing in a hin-walled, hyaline esicle o cha ac e is ic shape (Figu es 2F, 2G). Conidia a e cylind ical, ounded a bo h ends, s aigh , hyaline, (0)–1-sep a e, [(9–)11–13(–15)×2–4 μm], some imes         H I F G A B C E D Figu e 2. A, Conidiopho es o I. li iodend i; B, Mac o- and mic oconidia o Ilyonec ia li iodend i; C, Chlamydospo es in chains o “Cylind oca pon” paucisep a um; D, Conidiopho es o Campyl. ascicula e; E, Mac oconidia o Campyloca pon as- cicula e; F, Penicilla e conidiopho es o Cylind ocladiella pa a; G, Te minal esicles o Cyl. pa a; H, Conidia o Cyl. pa a; H, Chlamydospo es in chains o Cyl. pa a. Scale ba s: a‒c, ‒i = 10 μm; d‒e = 25 μm. 253 Vol. 52, No. 2, Augus , 2013 An upda e o black- oo disease o g ape ine becoming swollen a one end wi h age (Figu e 2H). Chlamydospo es a e abundan o mode a e, mo e equen ly a anged in chains han clus e s (Fig- u e 2I). Ae ial mycelium anges om da k o ligh b own (Figu es 3K, 3L) (C ous and Wing ield, 1993; Lomba d e al., 2012). Epidemiology Campyloca pon, “Cylind oca pon”, Cylind oclad- iella and Ilyonec ia species a e gene ally ega ded as pa hogens and/o sap obes o a wide ange o angio- spe m and gymnospe m hos s and subs a es in em- pe a e, sub- opical and opical egions wo ldwide (Vic o e al., 1998; Cha e i e al., 2011; Lomba d e al., 2012). In addi ion o g ape ine, hey ha e also been associa ed wi h oo o diseases o o he economi- cally impo an hos s (Cha e i e al., 2011; Lomba d e al., 2012), such as: Ac inidia chinensis Planch. (E - pe e al., 2011), Li iodend on ulipi e a L. (MacDonald and Bu le , 1981), Olea eu opaea L. (Ú bez-To es e al., 2012), Panax quinque olius L. (Rahman and Punja, 2005), Pe sea ame icana Mill. (Vi ale e al., 2012), Pinus adia a D. Don (Agus í-B isach e al., 2011b) o Pinus syl es is L. (Menkis and Bu okiene, 2012). Lomba d e al. (2013) ha e jus epo ed black oo o disease associa ed wi h he cul i a ion o P o eaceae cu low- e s in Sou h A ica, and desc ibed ou new Ilyonec-   A D C I E F G H L K J B Figu e 3. Colonies o black- oo pa hogens g own on PDA. A, “Cylind oca pon” des uc ans (CBS 301.93); B, “C.” paucisep- a um; C, “C.” ob usispo um; D, Ilyonec ia alcace ensis; E, I. li iodend i; F, I. no ozelandica; G, I. mac odidyma; H, I. o esensis; I, Campyloca pon ascicula e; J, Campyl. pseudo ascicula e; K, Cylind ocladiella pa a; L, Cyl. pe u iana. Phy opa hologia Medi e anea 260 C. Agus í-B isach and J. A mengol d ocladiella, and a compa ison wi h mo phologically simi- la gene a. Mycological Resea ch 97, 433–448. C ous P.W., A.J.L. Phillips and M.J. Wing ield, 1993. New e- co ds o Cylind ocladium and Cylind ocladiella spp. in Sou h A ica. Mycological Resea ch 42, 302–305. De F ancisco M.T., L. Ma in, R. Cobos, P. Ga cía-Bena ides and M.T. Ma in, 2009. 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