Checklist of the lichenicolous fungi of Greenland Prepared for the ...

Checklist of the lichenicolous fungi of Greenland 

Prepared for the NLF excursion to Greenland 2005 by Vagn Alstrup. The host lichens 

known from Greenland are mentioned, in brackets also hosts from outside Greenland. 

Localities etc. as in lichen list. 

* Previously known from the excursion area. 

** New to the area. 

*Abrothallus bertianus De Not. Parmelia fraudans, Nephroma parile, Pseudephebe 

pubescens (Melanelia, Neofuscelia, Cetraria) 

*Abrothallus parmeliarum (Sommerf.) Arnold, Parmelia spp. 

**Abrothallus welwitschii Mont. Ex Tul. 5, on Nephroma 

Acanthonitschkea peltigericola (Alstrup & Olech) O. E. Eriksson & R. Sant., (as 

Capronia peltigerae), Peltigera spp. 

Adelococcus alpestris (Zopf) Theiss. & Sydow, Acarospora glaucocarpa (brown spp.) 

Ahlesia lichenicola Fuckel, Lepraria neglecta (several hosts) 

Arctosporobolus alpinus (R. Sant., Alstrup & D. Hawksw.) Alstrup comb. ined., 

Ochrolechia, Pertusaria spp., Cladonia borealis. 1-12. Common. 

Arctosporobolus bryoriicolus Alstrup ined., Bryoria chalybeiformis 

Arthonia aspiciliae Alstrup & E. S. Hansen, Aspicilia elevata 

Arthonia clemens (Tul.) Th. Fr., Rhizoplaca peltata, Lecanora hageni, Protoparmelia 

badia 

Arthonia destruens Rehm, Lecanora leptacina (Xanthoria) 

Arthonia epicladonia (Nyl.) Alstrup & Zhurb., Cladonia subgen. Cladonia 

Arthonia epimela Norman, Biatora vernalis (Buellia punctata) 

*Arthonia epiphyscia Nyl., Physcia spp., Phaeorrhiza nimbosa (Phaeophyscia) 

*Arthonia excentrica Th. Fr., Leprocaulon 

?Arthonia fusca (Massal.) Hepp, several lichens 

*Arthonia fuscopurpurea (Tul.) R. Sant., Peltigera, Solorina, Psoroma hypnorum 

Arthonia galactinaria Leight., Lecanora dispersa-group 

Arthonia glacialis Alstrup & E. S. Hansen, Rhizoplaca melanophthalma 

Arthonia hymeniicola Heidmarsson & Alstrup ined., Xanthoria elegans (Caloplaca 

lithophila) 

Arthonia intexta Almq., Lecidella spp. 

Arthonia molendoi (Frauenf.) R. Sant., Xanthoria elegans, Caloplaca saxicola 

Arthonia obscurior Triebel, Pilophorus dovrensis 

Arthonia peltigerea Th. Fr., Peltigera, Solorina saccata, S. bispora 

Arthonia peltigerina (Almq.) H. Olivier, Peltigera, Solorina crocea 

*Arthonia pelvetii (Hepp) Almq., Peltigera, Cladonia carneola?, Pannaria pezizoides? 

Arthonia phaeophysciae Grube & Matzer, Phaeophyscia 

Arthonia stereocaulina (Ohlert) R. Sant. (as S. nephromiaria), Stereocaulon spp., 

Dactylina ramulosa? Leprocaulon subalbicans? 

Arthonia sublapponicae Alstrup, ined., Aspicilia sublapponica 

Arthonia tribeliae Zhurb., Dactylina arctica 

Arthonia varians (Davies) Nyl. (L. glaucomaria) Lecanora rupicola-group 

*Ascochyta santessonii Alstrup & D. Hawksw., Phaeophyscia sciastra 

Bacidia killiasii (Hepp) D. Hawksw., Peltigera 

Buellia adjuncta Th. Fr., Lecanora straminea, Rinodina olivaceobrunnea 

Buellia mylius-erichsenii Alstrup ined., Acarospora picea

*Buellia nialis (Bagl. & Car.) Hafellner, Xanthoria elegans, Caloplaca trachyphylla, 

Lecanora subradiosa? 

*Buellia pulverulenta (Anzi) Jatta, Physconia muscigena, Phaeophyscia sciastra, 

Physcia, Lecanora polytropa? 

Buellia thuleana Alstrup ined., Phaeophyscia sciastra, Rinodina sp. 

Carbonea aggregantula (Müll. Arg.) Diederich & Triebel, Lecanora polytropa 

Carbonea supersparsa (Nyl.) Hertel (Lecidea diexcipula), Lecanora polytropa, L. 

intricata (Rhizoplaca subdiscrepans) 

*Carbonea vitellinaria (Nyl.) Hertel, Candelariella spp., Pyrenopsis sp.? 

Cecidonia umbonella (Nyl.) Triebel & Rambold, Lecidea lapicida-group 

Cercidospora cecidiiformans Grube & Hafellner, Rhizocarpon, yellow spp. 

Cercidospora cephalodiorum Triebel & Grube, Pilophorus dovrensis 

Cercidospora epicarphinea (Nyl.) Grube & Hafellner (C. caudata), Xanthoria elegans 

(Caloplaca) 

*Cercidospora epipolytropa (Mudd) Arnold, Lecanora polytropa, Rhizoplaca 

melanophthalma. 1-12, very common. 

Cercidospora fatalis Alstrup ined., Xanthoria elegans 

Cercidospora groenlandica Alstrup ined., Ochrolechia upsaliensis 

Cercidospora lichenicola (Zopf) Hafellner, Solorina crocea, Protopannaria pezizoides 

( ) 

Cercidospora macrospora (Uloth) Hafellner & Nav.-Ros. (C. ulothii), 

Protoparmeliopsis, Rhizoplaca melanophthalma 

Cercidospora punctillata (Nyl.) R. Sant., (C. lichenicola) Solorina (Peltigera, 

Cladonia?). 

Cercidospora soror Obermayer & Triebel, Arthrorhaphis 

Cercidospora stereocaulorum (Arnold) Hafellner, Stereocaulon 

*Cercidospora trypetheliza (Nyl.) Hafellner & Obermayer, Arthrorhaphis alpina 

Cercidospora ullrichii Alstrup ined., Ochrolechia upsaliensis 

*Clypeococcum grossum (Körber) D. Hawksw., Umbilicaria vellea, U. cylindrica 

Corticifraga fuckelii (Rehm) D. Hawksw. & R. Sant., Peltigera 

Corticifraga peltigerae (Nyl.) D. Hawksw. & R. Sant., Peltigera 

Dacampia engeliana (Saut.) A. Massal., Solorina saccata 

Dacampia hookeri (Borrer) A. Massal. Solorina saccata, S. octospora 

Dactylospora allantoidea Alstrup & D. Hawksw., Neofuscelia pulla, Dimelaena oreina 

Dactylospora amygdalariae Triebel, Amygdalaria 

*Dactylospora aspiciliicola Alstrup & D. Hawksw., Koerberiella wimmeriana 

Dactylospora parasitica (Flörke) Zopf, Biatora cuprea (Ochrolechia, Pertusaria, 

Megaspora) 

Dactylospora purpurascens Triebel, Amygdalaria (Pilophorus dovrensis, Porpidea) 

Dactylospora rimicola (Müll. Arg.) Hafellner 

Dactylospora rinodinicola Alstrup & D. Hawksw., Rinodina turfacea 

Dactylospora saxatilis (Schaerer) Hafellner, Pertusaria (Ochrolechia) 

Dactylospora urceolata (th. Fr.) Arnold. Biatora vernalis, Lopadium coralloideum, 

Psoroma hypnorum (several lichens) 

Dahawksia hauganii Alstrup & Zhurb. ined., Ochrolechia lactea 

Deichmannia verrucispora Alstrup & D. Hawksw., Diploschistes muscorum 

*Didymella sphinctrinoides (Zwackh) Berl. & Vogl., Physcia magnussonii 

Diplotomma alboatrum (Hoffm.) Flot. (Buellia nivalis), Caloplaca trachyphylla, 

Xanthoria elegans, Lecanora subradiosa? 

Echinothecium reticulatum Zopf, Parmelia

Endococcus fusiger Th. Fr. & Almq., Rhizocarpon, brown and grey spp. 

Endococcus latisporus Alstrup ined., Aspicilia sublapponica, Aspicilia sp. 

Endococcus macrosporus (Arnold) Nyl., Rhizocarpon, yellow spp. 

Endococcus parietinus (Lindsay) Clauz. & Cl. Roux, Caloplaca cerina (Xanthoria 

parietina) 

Endococcus perpusillus Nyl., Rhizocarpon spp., Schaereria fuscocinerea 

*Endococcus propinquus (Körber) D. Hawksw., Rhizocarpon sp., Porpidia, Lecanora 

intricata (crustose, saxicolous lichens, esp. Porpidia) 

*Endococcus rugulosus Nyl. Koerberiella wimmeriana, Aspicilia, Protoparmelia 

(crustose, saxicolous lichens) 

?Endococcus stigma (Körber) Stizenb. Acarospora 

Endococcus tetrasporus Alstrup ined., Candelariella 

Epilichen glauconigellus (Nyl.) Hafellner, Baeomyces 

*Epilichen scabrosus (Ach.) Clem, Baeomyces 

*Everniicola flexispora D. Hawksw., Nephroma arcticum (Evernia prunastri) 

Fayodia striatula (Kühner) Singer, Peltigera 

Geltingia associata (Th. Fr.) Alstrup & D. Hawksw., Pertusaria dactylina 

(Ochrolechia, Pertusaria) 

Geltingiomyces diskoensis Alstrup ined., Aspicilia annulata 

**Graphium aphthosae Alstrup & D. Hawksw., Peltigera aphthosa, Peltigera spp., 

Solorina bispora. 9, 10 on P. aphthosa. 

Hainesia cetrariae Alstrup & A. Larsen ined., Cetraria islandica 

*Homostegia piggottii (Berk. & Broome) P. Karst., Parmelia 

Hymenobia aporea (Nyl.) D. Hawksw., Lecidea 

*Illosporium carneum Fr., Peltigera didactyla, Peltigera spp. 

Intralichen christiansenii (D. Hawksw.) D. Hawksw. & M. S. Cole, Arthonia 

exentrica, Rinodina turfacea, Coriscium viride?, Stereocaulon condensatum? (several 

licens) 

Intralichen diploschistei Alstrup ined., Diploschistes muscorum 

*Intralichen lichenicola (M. S. Christ. & D. Hawksw.) D. Hawksw. & M. S. Cole, 

Candelariella spp. Lecanora polytropa, L. fuscescens (many hosts) 

*Intralichen lichenum (Diederich) D. Hawksw, & M. S. Cole, Protoblastenia 

incrustans, Ionaspis epulotica, Lecanora polytropa 

*Kalaallia reactiva Alstrup & D. Hawksw., Ionaspis lacustris 

Karschia alpicolae Alstrup & D. Hawksw., Allantoparmelia alpicola 

Karsteniomyces tuberculosis Alstrup & D. Hawksw., Peltigera 

Lasiosphaeriopsis christiansenii Alstrup & D. Hawksw., Porpidia tuberculosa 

*Lasiosphaeriopsis stereocaulicola (Lindsay) O. E. Eriksson & R. Sant., Stereocaulon 

Leptosphaerulina peltigerae (Fuckel) Riedl, Peltigera 

Lecidea hymeneliicola Alstrup & D. Hawksw., Ionaspis lacustris 

*Lecidella lecanoricola Alstrup, D. Hawksw. & R. Sant., Lecanora cenisea 

Lichenochora constrictella (Müll. Arg.) Hafellner, Fulgensia bracteata 

Lichenoconium lecanorae (Jaap) D. Hawksw, Rhizoplaca melanophthalma (many 

hosts) 

Lichenoconium usneae (Anzi) D. Hawksw., Parmelia saxatilis, Abrothallus 

parmeliarum (many hosts) 

Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw., Caloplaca alcarum, 

Rhizoplaca, Pertusaria, Xanthoria elegans (many hosts) 

Lichenopeltella cetrariicola (Nyl.) R. Sant., Cetraria spp. 

*Lichenopeltella cladoniarum E. S. Hansen & Alstrup, Cladonia subgen. Cladina

*Lichenopuccinia poeltii D. Hawksw. & Hafellner, Parmelia 

*Lichenosticta alcicornaria (Lindsay) D. Hawksw., Cladonia 

*Lichenostigma arctoparmeliae R. Sant. ined. Arctoparmelia incurva 

*Lichenostigma cosmopolites , Xanthoparmelia conspersa 

Lichenostigma rugosum Thor, Diplosctistes 

Lichenostigma semiimmersa Hafellner, Buellia elegans 

*Llimoniella neglecta (Vainio) Triebel & Rambold, Lepraria neglecta 

**Marchandiomyces corallinus (Roberge) Diederich & D. Hawksw., Parmelia, 

Pannanaria, Melanelia hepatizon (many hosts). 9, 10, 12. 

Merismatium coccisporum (Norman) Vouaux, Rhizocarpon etc. 

*Merismatium decolorans (Arnold) Triebel, Arthrorhaphis citrinella 

Merismatium heterophractum (Nyl.) Vouaux, Nephroma parile (several hosts) 

*Merismatium nigritellum (Nyl.) Vouaux (M. lopadii), Euopsis macrocarpa, Lepraria 

sp. (several hosts) 

Merismatium thamnoliicola Alstrup & E. S. Hansen, Thamnolia 

*Muellerella lichenicola (Sommerf.) Fr. D. Hawksw., saxicolous, crustose lichens 

*Muellerella pygmaea (Körber) D. Hawksw., saxicolous, crustose lichens 

Neolamya peltigerae (Mont.) Theissen & Sydow, Peltigera 

Niesslia cladoniicola D. Hawksw. & Gams, Cladonia subgen. Cladina 

Niesslia kununguakii Alstrup & E. S. Hansen, Arthrorhaphis alpina 

Nigropuncta groenlandica Alstrup, unidentified host 

Opegrapha glaucomaria (Nyl.) Källsten, Lecanora rupicola 

Opegrapha stereocaulicola Alstrup & D. Hawksw., Stereocaulon alpinum 

(Sterocaulon spp.) 

*Phacopsis oxyspora (Tul.) Triebel & Rambold, Parmelia, parmeliaceous lichens 

Phacopsis oxyspora var. defecta Triebel & Rambold, Parmelia saxatilis, P. fraudans 

(Platismatia, Melanelia) 

Phaeopyxis muscigenae Alstrup & E. S. Hansen, Physconia muscigena 

Phaeospora arctica Horáková & Alstrup (P. rimosicola in Alstrup & Hawksworth 

1990), Arctoparmelia centrifuga (Dactylina madreporiformis, Cetraria andrejevii) 

Phaeospora parasitica (Lönnr.) Arnold? 

Phaeospora verrucariae Alstrup & E. S. Hansen, Verrucaria placynthiae. 

Phaeosporobolus umbilicariae Alstrup ined., Umbilicaria rigida 

Phaeosporobolus usneae D. Hawksw. & Hafellner, Physcia caesia (mostly epiphytic 

macrolichens) 

*Phoma epiphyscia Vouaux, Phaeophyscia sciastra (Phaeophyscia orbicularis, 

Xanthoria parietina) 

*Phylliscum demangeonii (Moug. & Nestler) Nyl. Ochrolechia frigida (mostly 

lichenized) 

Polycoccum bryonthae (Arnold) Vezda?, Caloplaca sp.(Lecanora epibryon) 

Polycoccum microsticticum (Mudd) Arnold, Ionaspis lacustris, Rhizocarpon 

disporum (several crustose, saxicolous hosts) 

Polycoccum pulvinatum (Eitner) R. Sant., Physcia 

Polycoccum sporastitiae (Anzi) Arnold, Sporastatia 

Polycoccum squamaroides (Mudd) Arnold, Pertusaria, Placopsis 

*Polycoccum trypethelioides (Th. Fr.) R. Sant., Stereocaulon 

Polycoccum vermicularium (Lindsay) D. Hawksw., Thamnolia 

Polydesmia lichenis Huhtinen & R. Sant., Peltigera 

Polysporina lapponica (Schaerer) Degel., Acarospora, brown spp. 

Pronectria erythrinalla (Nyl.) Lowen, Peltigera

Pronectria ornamentata (D. Hawksw.) Lowen, Peltigera 

Pronectria robergei (Mont. & Desm.) Lowen, Peltigera 

Protothelenella croceae (Bagl. & Carestia) Hafellner & H. Mayrhofer, Peltigera 

didactyla (Solorina croceae) 

Protothelenella santessonii Mayrhofer, Cladonia 

Pyrenidium actinellum Nyl.?, ? (many hosts) 

Pyrenidium hyalosporum Alstrup, D. Hawksw. & R. Sant., Placopsis gelida 

Raciborskiomyces peltigericola (D. Hawksw.) M. E. Barr, Peltigera 

Refractohilum galligenum D. Hawksw., Nephroma spp. 

Refractohilum peltigerae (Keissler) D. Hawksw., Peltigera 

*Rhagadostoma lichenicola (De Not.) Keissler, Solorina crocea, Peltigera rufescens? 

9,10,12. 

Rhexophiale parasitica (Alstrup & E. S. Hansen) Alstrup comb. ined., Hymenelia 

cyanocarpa 

*Rhizocarpon destructans Alstrup, Lecanora polytropa 

*Rhizocarpon narssaqensis Alstrup, Lecidea atromarginata 

Rhymbocarpus neglectus (Vainio) Diederich & Etayo, Lepraria sp., Leprocaulon 

subalbicans 

Rhymbocarpus stereocaulorum (Alstrup & D. Hawksw.) Etayo & Diederich, 

Stereocaulon 

Rinodina egedeana (Lindsay) Alstrup & D. Hawksw., Peltigera sp., Parmelia saxatilis, 

Cladonia pocillum. 

Rinodina turfacea (Wahlenb) Körber, Peltigera (mostly lichenized). 

Roselliniella cladoniae (Anzi) Matzer & Hafellner, Cladonia 

Roselliniopsis gelidaria (Mudd) Matzer (Polycoccum gelidarium), Placopsis gelida 

*Roselliniopsis ventosa (Rostrup) Alstrup, Placopsis gelida, Buellia chionea 

*Rosellinula frustulosae (Vouaux) R. Sant., Schistoplaca argopholis. 5. 

*Sagediopsis barbara (Th. Fr.) R. Sant. & Triebel, Porpidia glaucophaea 

*Sagediopsis campsteriana (Lindsay) D. Hawksw. & R. Sant., Ochrolechia 

Sclerococcum simplex D. Hawksw., Melanelia substygia (Pertusaria) 

*Sclerococcum sphaerale (Ach.) Th. Fr., Pertusaria 

Scutula cladoniicola Alstrup & D. Hawksw., Cladonia stricta (Cladonia subgen. 

Cladonia) 

Scutula epiblastemica (Wallr.) Rehm, Peltigera 

Scutula krempelhuberi Körber, Solorina 

Scutula miliaris (Wallr.) Trevisan, Peltigera 

*Scutula solorinaria (Nyl.) P. Karsten, Solorina bispora 

*Scutula stereocaulorum (Anzi) Körber, Stereocaulon 

Scutula tuberculosa (Th. Fr.) Rehm; Peltigera, Solorina crocea 

Skyttea arenicola Alstrup & E. S. Hansen, Cladonia pocillum? 

Skyttea fusispora Sherwood, D. Hawksw. & Coppins, ? 

Skyttella muelleri (Willey) D. Hawksw. & R. Sant., Peltigera 

*Sphaerellothecium araneosum (Rehm & Arnold) Zopf, Ochrolechia, Pertusaria, 

Arctoparmelia?, Calvitimela armeniaca? 1-12, very common on Ochrolechia frigida. 

*Sphaerellothecium araneosum var. cladoniae Zhurb. & Alstrup, Cladonia subgen. 

Cladonia 

Sphaerellothecium atrynea (Arnold) Roux & Triebel 

*Sphaerellothecium cladoniicola E. S. Hansen & Alstrup, Cladonia subgen. Cladina 

Sphaerellothecium melaneliicola Alstrup ined., Melanelia 

Sphaerellothecium minutum Hafellner, Sphaerophorus

Sphaerulina intermedia 

Steinia geophana Nyl.) B. Stein, Peltigera (algicolous?) 

*Stigmidium conspurcans (Th. Fr.) Triebel & R. Sant., Psora rubiformis 

Stigmidium ephebes (Henssen) D. Hawksw., Ephebe 

Stigmidium frigidum (Sacc.) Alstrup & D. Hawksw., Thamnolia 

Stigmidium glebarum (Arnold) Hafellner, Toninia sedifolia 

Stigmidium marinum (Deakin) Swinscow, marine Verrucaria spp. 

*Stigmidium microcarpum Alstrup & J. C. Davis, Flavocetraria cucullata. 5, 10. 

Stigmidium peltideae (Vainio) R. Sant., Peltigera 

Stigmidium psorae (Anzi) Hafellner, Psora 

Stigmidium solorinarium (Vainio) D. Hawksw., Solorina saccata 

Stigmidium stygnospilum (Minks) R. Sant.? Catrapyrenium lachneum 

Stigmidium tabacina (Arnold) Triebel, Toninia tristis 

Stigmidium spp., Psoroma hypnorum, Melanelia substygia 

Syzogospora bachmannii Diederich & M. S. Christ, Cladonia 

Taeniolella christiansenii Alstrup & D. Hawksw. Stereocaulon alpinum, Arthonia 

stereocaulina 

Taeniolella delicata M. S. Christ. & D. Hawksw., Icmadophila ericetorum (several 

hosts) 

Taeniolella ionaspisicola Alstrup & E. S. Hansen, Ionaspis odora 

Taeniolella pertusariicola D. Hawksw. & H. Mayrhofer, Pertusaria spp. 

Taeniolella strictae Alstrup, Cladonia stricta 

Thamnogalla crombiei (Mudd)D. Hawksw., Thamnolia vermicularis 

Thelocarpon epibolum Nyl., Peltigera, Solorina crocea 

Tomentella fuscella Lee coll., Solorina saccata, Peltigera sp. 

Unguiculariopsis cribriformis (Norman) Alstrup & D. Hawksw., Pertusaria 

(Ochrolechia) 

*Unguiculariopsis groenlandiae (Alstrup & D. Hawksw.) Etayo & Diederich, 

Caloplaca citrina (Fulgensia) 

*Vouauxiella caloplacae Alstrup, Caloplaca scopularis 

*Vouauxiomyces santessonii D. Hawksw., Parmelia saxatilis 

Weddellomyces tartariicola (Lindsay) Alstrup & D. Hawksw., Ochrolechia frigida 

Zwackhiomyces sphinctrinoides (Zwackh) Grube & Hafellner, Lecanora albescens, L. 

campestris 

References 

Alstrup, V. 1981. Notes on some lichens and lichenicolous fungi from Greenland. 

Nord. J. Bot. 1:120-124. 

Alstrup, V. 1993. New on lichens and lichenicolous fungi from the Nordic countries. 

Graphis Scripta 5: 96-104. 

Alstrup, V. 2004. New records of distribution of lichens and lichenicolous fungi. 

Graphis Scripta 16: 46-57. 

Alstrup, V. & Hansen, E. S. 2001. New lichens and lichenicolous fungi from 

Greenland. Graphis Scripta 12: 41-50. 

Alstrup, V. & Hawksworth 1990. The lichenicolous fungi of Greenland. Meddelelser 

om Grønland. Bioscience 31. 

Alstrup, V. et al. (in press). The lichenicolous fungi of the Skibotn area, Troms 

County, Norway. 

Hansen, E. S. 1998. Notes on some new and interesting lichens and lichenicolous

fungi from Greenland. Graphis Scripta 9: 7-9. 

Hansen, E. S. & Alstrup, V. 1995. The lichenicolous fungi on Cladonia subgen. 

Cladina in Greenland. Graphis Scripta 7: 33-38. 

Hansen, E. S. & Lund, P. M. 2003. The lichen flora and reindeer grazing in the 

Isortoq area, South West Greenland. Folia Cryptogamica Estonica 40: 7-14. 

Hansen, E. S. & Obermayer, W. 1999. Notes on Arthrorhaphis and its lichenicolous 

fungi in Greenland. Bryologist 102: 104-107. 

Horáková, J. & Alstrup, V. Phaeospora arctica sp. nov. Graphis Scripta 

Kümmerling, H., Triebel, D. & Rambold, G. 1993. Lepraria neglecta and its 

lichenicolous fungi. Bibl. Lichenol. 53: 147-160. 

Triebel, D. 1989. Lecideicole Ascomyceten. Bibliotheca Lichenologica 35. 

Zhurbenko, M. & Alstrup, V. 2004. Lichenicolous fungi on Cladonia mainly from the 

Arctic. Symb. Bot. Ups- 34: 477-499.

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