In this paper we present an updated checklist of thallus-forming Laboulbeniomycetes (Ascomycota, Pezizomycotina), that is, the orders Herpomycetales and Laboulbeniales, from Belgium and the Netherlands. Two species are newly described based on morphology, molecular data (ITS, LSU ribosomal DNA) and ecology (host association). These are Hesperomyces halyziae on Halyzia sedecimguttata (Coleoptera, Coccinellidae) from both countries and Laboulbenia quarantenae on Bembidion biguttatum (Coleoptera, Carabidae) from Belgium. In addition, nine new country records are presented. For Belgium: Laboulbenia aubryi on Amara aranea (Coleoptera, Carabidae) and Rhachomyces spinosus on Syntomus foveatus (Coleoptera, Carabidae). For the Netherlands: Chitonomyces melanurus on Laccophilus minutus (Coleoptera, Dytiscidae), Euphoriomyces agathidii on Agathidium laevigatum (Coleoptera, Leiodidae), Laboulbenia fasciculata on Omophron limbatum (Coleoptera, Carabidae), Laboulbenia metableti on Syntomus foveatus and S. truncatellus (Coleoptera, Carabidae), Laboulbenia pseudomasei on Pterostichus melanarius (Coleoptera, Carabidae), Rhachomyces canariensis on Trechus obtusus (Coleoptera, Carabidae), and Stigmatomyces hydrelliae on Hydrellia albilabris (Diptera, Ephydridae). Finally, an identification key to 140 species of thallus-forming Laboulbeniomycetes in Belgium and the Netherlands is provided. Based on the combined data, we are able to identify mutual gaps that need to be filled as well as weigh the impact of chosen strategies (fieldwork, museum collections) and techniques in these neighboring countries. The aim of this work is to serve as a reference for studying Laboulbeniomycetes fungi in Europe.

Ghent University Ghent Belgium Ghent University Ghent Belgium

Meise Botanic Garden Meise Belgium Meise Botanic Garden Meise Belgium

Purdue University West Lafayette Indiana United States of America Purdue University West Lafayette United States of America

University of South Bohemia České Budějovice Czech Republic University of South Bohemia Budějovice Czech Republic

Zobrazit více v PubMed

Arndt E, Santamaría S. (2004) Laboulbeniales (Ascomycota) of the Canary Islands. Vieraea 32: 107–115.

Bánhegyi J. (1960) Contributions à la connaissance des Laboulbéniales de la péninsule des Balkans. Annales Universitatis Scientiarum Budapestiensis de Rolando Eötvös Nominatae; Sectio Biológica 3: 49–67.

Baral HO, Weber E, Marson G, Quijada L. (2018) A new connection between wood saprobism and beetle endosymbiosis: the rarely reported saprobic discomycete Tromeropsis is congeneric with the symbiotic yeast Symbiotaphrina (Symbiotaphrinales, Xylonomycetes) and two asexual morphs misplaced in Hyphozyma. Mycological Progress 17: 215–254. 10.1007/s11557-017-1340-y DOI

Beccaloni GW, Eades DC, Hopkins H. (2014) Cockroach species file online. Version 5.0/5.0. http://cockroach.speciesfile.org [accessed 7 July 2020]

Benjamin RK. (1971) Introduction and supplement to Roland Thaxter’s contribution towards a monograph of the Laboulbeniaceae. Bibliotheca Mycologica 80: 1–155.

Bennett C, Aime MC, Newcombe G. (2011) Molecular and pathogenic variation within Melampsora on Salix in western North America reveals numerous cryptic species. Mycologia 103: 1004–1018. 10.3852/10-289 PubMed DOI

Blackwell M. (1994) Minute mycological mysteries: the influence of arthropods on the lives of fungi. Mycologia 86: 1–17. 10.1080/00275514.1994.12026371 DOI

Blackwell M. (2011) The Fungi: 1, 2, 3 … 5.1 million species? American Journal of Botany 98: 426–438. 10.3732/ajb.1000298 PubMed DOI

Blackwell M, Haelewaters D, Pfister DH. (2020) Laboulbeniomycetes: Evolution, natural history, and Thaxter’s final word. Mycologia: 1–12. 10.1080/00275514.2020.1718442 PubMed DOI

Boedijn K. (1923) On the development of Stigmatomyces. Mededelingen van de Nederlandse Mycologische Vereniging 13: 91–97.

Capella-Gutiérrez S, Silla-Martínez JM, Gabaldón T. (2009) TrimAl: a tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25: 1972–1973. 10.1093/bioinformatics/btp348 PubMed DOI PMC

Cavara F. (1899) Di una nuova Laboulbeniaceae: Rickia wasmannii, nov. gen. et nov. spec. Malpighia 13: 173–188.

Chernomor O, Von Haeseler A, Minh BQ. (2016) Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology 65: 997–1008. 10.1093/sysbio/syw037 PubMed DOI PMC

Collart A. (1945) A propos des Laboulbéniacées. Bulletin Mensuel des Naturalistes Belges 26: 98–103.

Collart A. (1947) A la découverte des Laboulbéniales. Bulletin et Annales de la Société Entomologique de Belgique 83: 21–35.

Cooke MC. (1892) Vegetable Wasps and Plant Worms: A Popular History of Entomogenous Fungi, or Fungi Parasitic Upon Insects. Society for Promoting Christian Knowledge, London, 364 pp 10.5962/bhl.title.34922 DOI

Darriba D, Taboada GL, Doallo R, Posada D. (2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9: 1–772. 10.1038/nmeth.2109 PubMed DOI PMC

De Kesel A. (1994) Phaulomyces simplocariae sp. nov. (Ascomycetes, Laboulbeniales) from Simplocaria semistriata (Coleoptera, Byrrhidae). Mycotaxon 50: 191–198.

De Kesel A. (1998) Identification and host-range of the genus Laboulbenia in Belgium. Sterbeeckia 18: 13–31.

De Kesel A. (1999) Peyritschiella heinemanniana sp. nov. (Ascomycetes, Laboulbeniales) a new ectoparasitic species from Xantholinus longiventris (Coleoptera, Staphylinidae). Belgian Journal of Botany 131: 176–180.

De Kesel A. (2010) Laboulbeniales (Ascomycetes) from coprophilic Coleoptera on Galloway cow dung in Belgium. Polish Botanical Journal 55: 353–358.

De Kesel A, Gerstmans C. (2012) Distolomyces forficulae (Laboulbeniales), a common parasite on Forficula auricularia (Dermaptera). Sterbeeckia 31: 43–44.

De Kesel A, Gerstmans C, Haelewaters D. (2020) Catalogue of the Laboulbeniomycetes of Belgium. Sterbeeckia 36: 3–143.

De Kesel A, Haelewaters D. (2014) Laboulbenia slackensis and L. littoralis sp. nov. (Ascomycota, Laboulbeniales), two sibling species as a result of ecological speciation. Mycologia 106: 407–414. 10.3852/13-348 PubMed DOI

De Kesel A, Haelewaters D. (2019) Laboulbeniales (Fungi, Ascomycota) of cholevine beetles (Coleoptera, Leiodidae) in Belgium and the Netherlands. Sterbeeckia 35: 60–66.

De Kesel A, Haelewaters D, Dekoninck W. (2016) Myrmecophilous Laboulbeniales (Ascomycota) in Belgium. Sterbeeckia 34: 3–6.

De Kesel A, Haelewaters D, Gerstmans C. (2013) Two interesting species of Rickia (Laboulbeniales) from coastal habitats in Belgium and the Netherlands. Sterbeeckia 32: 6–10.

De Kesel A, Rammeloo J. (1992) Checklist of Laboulbeniales (Ascomycetes) of Belgium. Belgian Journal of Botany 124: 204–214.

De Kesel A, Werbrouck T. (2008) Belgian records of Laboulbeniales from aquatic insects. Sterbeeckia 28: 48–54.

De Weggheleire S. (2019) Biodiversity of the genus Laboulbenia: are there cryptic species? M.Sc. Thesis, University of Antwerp, Belgium.

Drummond AJ, Suchard MA, Xie D, Rambaut A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecoluar Biology and Evolution 29: 1969–1973. 10.1093/molbev/mss075 PubMed DOI PMC

Edgar RC. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792–1797. 10.1093/nar/gkh340 PubMed DOI PMC

Ehrlich PR, Raven PA. (1964) Butterflies and plants: A study in coevolution. Evolution 18: 586–608. 10.1111/j.1558-5646.1964.tb01674.x DOI

Gardes M, Bruns TD. (1993) ITS primers with enhanced specificity for basidiomycetes – application to the identification of mycorrhizae and rusts. Molecular Ecology 2: 113–118. 10.1111/j.1365-294X.1993.tb00005.x PubMed DOI

Gernhard T. (2008) The conditioned reconstructed process. Journal of Theoretical Biology 253: 769–778. 10.1016/j.jtbi.2008.04.005 PubMed DOI

Goldmann L, Weir A, Rossi W. (2013) Molecular analysis reveals two new dimorphic species of Hesperomyces (Ascomycota, Laboulbeniomycetes) parasitic on the ladybird Coleomegilla maculata (Coleoptera, Coccinellidae). Fungal Biology 117: 807–813. 10.1016/j.funbio.2013.10.004 PubMed DOI

Grünig CR, Duo A, Sieber TN, Holdenrieder O. (2008) Assignment of species rank to six reproductively isolated cryptic species of the Phialocephala fortinii s.l.–Acephala applanata species complex. Mycologia 100: 47–67. 10.3852/mycologia.100.1.47 PubMed DOI

Gutierrez AC, Ordoqui E, Leclerque A, Lastra CL. (2020) A new species of Herpomyces (Laboulbeniomycetes: Herpomycetales) on Periplaneta fuliginosa (Blattodea: Blattidae) from Argentina. Mycologia: 1–8. 10.1080/00275514.2020.1726134 PubMed DOI

Haelewaters D. (2012) The first record of Laboulbeniales (Fungi, Ascomycota) on ants (Hymenoptera, Formicidae) in the Netherlands. Ascomycete. org 4: 65–69.

Haelewaters D. (2013) Nieuwe Laboulbeniales. In: Peeters T, van Eck A, Cramer T. (Eds) Natuurstudie in De Kaaistoep.Verslag 2012, 18e onderzoeksjaar. TWM Gronden BV, Natuurmuseum Brabant & KNNV-afdeling Tilburg, 23–26.

Haelewaters D, Dima B, Abdel-Hafiz BII, Abdel-Wahab MA, Abul-Ezz SR, Acar I, Aguirre-Acosta E, Aime MC, Aldemir S, Ali M, Ayala-Vásquez O, Bakhit MS, Bashir H, Battistin E, Bendiksen E, Castro-Rivera R, Çolak ÖF, De Kesel A, de la Fuente JI, Dizkırıcı A, Hussain S, Jansen GM, Kaygusuz O, Khalid AN, Khan J, Kiyashko AA, Larsson E, Martínez-González CR, Morozova OV, Niazi AR, Noordeloos ME, Pham THG, Popov ES, Psurtseva NV, Schoutteten N, She H, Türkekul İ, Verbeken A, Ahmad H, Afshan NS, Christe P, Fiaz M, Glaizot O, Liu J, Majeed J, Markotter W, Nagy A, Nawaz H, Papp V, Péter Á, Pfliegler WP, Qasim T, Riaz M, Sándor AD, Szentiványi T, Voglmayr H, Yousaf N, Krisai-Greilhuber I. (in press) Fungal Systematics and Evolution 6. Sydowia 72.

Haelewaters D, Boer P, Noordijk J. (2015a) Studies of Laboulbeniales (Fungi: Ascomycota) on Myrmica ants: Rickia wasmannii in the Netherlands. Journal of Hymenoptera Research 44: 39–47. 10.3897/JHR.44.4951 DOI

Haelewaters D, De Kesel A. (2013) A new species of Cantharomyces (Laboulbeniales, Ascomycota) from the Netherlands. Mycotaxon 123: 467–472. 10.5248/123.467 DOI

Haelewaters D, De Kesel A, Pfister DH. (2018) Integrative taxonomy reveals hidden species within a common fungal parasite of ladybirds. Scientific Reports 8: 15966. 10.1038/s41598-018-34319-5 PubMed DOI PMC

Haelewaters D, De Kesel A, Gorczak M, Bao K, Gort G, Zhao SY, Pfister DH. (2019a) Laboulbeniales (Ascomycota) of the Boston Harbor Islands II: species parasitizing Carabidae, and the Laboulbenia flagellata species complex. Northeastern Naturalist 25(Special Issue 9): 110–149. 10.1656/045.025.s906 DOI

Haelewaters D, De Kock G, van Wielink P. (2015b) Nieuwe Laboulbeniales in De Kaaistoep. In: Peeters T, van Eck A, Cramer T. (Eds) Natuurstudie in De Kaaistoep.Verslag 2014, 20e onderzoeksjaar. TWM Gronden BV, Natuurmuseum Brabant & KNNV-afdeling Tilburg, 11–18.

Haelewaters D, Gorczak M, Pfliegler WP, Tartally A, Tischer M, Wrzosek M, Pfister DH. (2015c) Bringing Laboulbeniales into the 21st century: enhanced techniques for extraction and PCR amplification of DNA from minute ectoparasitic fungi. IMA Fungus 6: 363–372. 10.5598/imafungus.2015.06.02.08 PubMed DOI PMC

Haelewaters D, Nuytinck J, De Kesel A. (2012a) Laboulbeniales in Nederland: een introductie. Natuurhistorisch Maandblad 101: 88–93.

Haelewaters D, Pfliegler WP, Gorczak M, Pfister DH. (2019b) Birth of an order: comprehensive molecular phylogenetic study reveals that Herpomyces (Fungi, Laboulbeniomycetes) is not part of Laboulbeniales. Molecular Phylogenetics and Evolution 133: 286–301. 10.1016/j.ympev.2019.01.007 PubMed DOI

Haelewaters D, van Wielink P. (2016) Hesperomyces virescens (Laboulbeniales) op een nieuwe gastheer. In: Peeters T, Cramer T, van Eck A. (Eds) Natuurstudie in De Kaaistoep.Verslag 2015, 21e onderzoeksjaar. TWM Gronden BV, Tilburg, 9–12.

Haelewaters D, van Wielink P, van Zuijlen JW, Verbeken A, De Kesel A. (2012b) New records of Laboulbeniales (Fungi, Ascomycota) for the Netherlands. Entomologische Berichten 72: 175–183.

Haelewaters D, Vorst O, De Kesel A. (2014) New and interesting Laboulbeniales (Fungi, Ascomycota) from the Netherlands. Nova Hedwigia 98: 113–125. 10.1127/0029-5035/2013/0150 DOI

Haelewaters D, Zhao SY, Clusella-Trullas S, Cottrell TE, De Kesel A, Fiedler L, Herz A, Hesketh H, Hui C, Kleespies RG, Losey JE, Minnaar IA, Murray KM, Nedvěd O, Pfliegler WP, Raak-van den Berg CL, Riddick EW, Shapiro-Ilan DI, Smyth RR, Steenberg T, van Wielink PS, Viglášová S, Zhao Z, Ceryngier P, Roy HE. (2017) Parasites of Harmonia axyridis: current research and perspectives. BioControl 62: 355–371. 10.1007/s10526-016-9766-8 DOI

Hawksworth DL, Lücking R. (2017) Fungal diversity revisited: 2.2 to 3.8 million species. Microbiology spectrum 5. 10.1128/microbiolspec.FUNK-0052-2016 PubMed DOI PMC

Hoang DT, Chernomor O, Von Haeseler A, Minh BQ, Vinh LS. (2017) UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–22. 10.1093/molbev/msx281 PubMed DOI PMC

Huldén L. (1983) Laboulbeniales (Ascomycetes) of Finland and adjacent parts of the U.S.S.R. Karstenia 23: 31–136. 10.29203/ka.1983.221 DOI

Index Fungorum (2020) Index Fungorum. http://www.indexfungorum.org [accessed 19 May 2020]

Justamond M. (2019) Halyzia 16-guttata https://www.flickr.com/photos/rock_wolf/49500355673/ [accessed 21 July 2020]

Kalyaanamoorthy K, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. 10.1038/nmeth.4285 PubMed DOI PMC

Kirk P. (2019) Catalogue of Life. http://www.catalogueoflife.org [accessed 14 July 2020]

Kossen WJ. (1936) Observations on a Dutch member of the Laboulbeniaceae. Annales Mycologici 34: 281–285.

Kossen WJ. (1938) Een goede vangst. De Levende Natuur 42: 152–154.

Lee Y-B, Na Y-H, Jang I-H, Lim C-K, Yang Y-K. (2007) Notes on three species of the genus Euphoriomyces (Laboulbeniales) from Korea. Mycobiology 35: 1–5. 10.4489/MYCO.2007.35.1.001 PubMed DOI PMC

Li YM, Shivas RG, Cai L. (2017) Cryptic diversity in Tranzscheliella spp. (Ustilaginales) is driven by host switches. Scientific Reports 7: 1–13. 10.1038/srep43549 PubMed DOI PMC

Liu J, Haelewaters D, Pfliegler WP, Page RA, Dick CW, Aime MC. (2020) A new species of Gloeandromyces from Ecuador and Panama revealed by morphology and phylogenetic reconstruction, with a discussion of secondary barcodes in Laboulbeniomycetes taxonomy. Mycologia. 10.1080/00275514.2020.1781496 PubMed DOI

Maire R. (1920) Troisième contribution à l’étude des Laboulbéniales de l’Afrique du Nord. Bulletin de la Societé d’Histoire Naturelle de l’Afrique du Nord 11: 123–138. [143–170.]

Majerus MEN, Williams Z. (1989) The distribution and life history of the orange ladybird, Halyzia sedecimguttata (L.) (Coleoptera: Coccinellidae) in Britain. Entomologist’s Gazette 40: 71–78.

Majewski T. (1994) The Laboulbeniales of Poland. Polish Botanical Studies 7: 3–466.

Majewski T. (1999) New and rare Laboulbenialis (Ascomycetes) from the Białowieża Forest (NE Poland). Acta Mycologica 34: 7–39. 10.5586/am.1999.001 DOI

Meijer J. (1975) Carabid (Coleoptera, Carabidae) migration studied with Laboulbeniales (Ascomycetes) as biological tags. Oecologia 19: 99–103. 10.1007/BF00369094 PubMed DOI

Miadlikowska J, McCune B, Lutzoni F. (2002) Pseudocyphellaria perpetua, a new lichen from Western North America. Bryologist 105: 1–10. 10.1639/0007-2745(2002)105[0001:PPANLF]2.0.CO;2 DOI

Middelhoek A. (1941) Dichomyces princeps Thaxter. Fungus 12: 56–57.

Middelhoek A. (1942) Een nieuwe Laboulbeniaceae voor ons land. Fungus 13: 52–53.

Middelhoek A. (1943a) Laboulbeniaceae in Nederland. Nederlands Kruidkundig Archief 53: 86–115.

Middelhoek A. (1943b) Parasitaire keverschimmels uit Zuid-Limburg. Natuurhistorisch Maandblad 32: 58–60.

Middelhoek A. (1943c) Enige nieuwe Laboulbeniales voor ons land. Fungus 14: 57–59.

Middelhoek A. (1943d) Enige nieuwe Laboulbeniales voor ons land (vervolg). Fungus 14: 71–72.

Middelhoek A. (1945) Twee keverschimmels op een gastheer. Fungus 16: 6–8.

Middelhoek A. (1947a) Laboulbeniaceae in Nederland II. Nederlands Kruidkundig Archief 54: 232–239.

Middelhoek A. (1947b) Wij en de keverschimmels. Natura 44: 89–93.

Middelhoek A. (1949) Laboulbeniaceae in Nederland III. Nederlands Kruidkundig Archief 56: 249–260.

Middelhoek A. (1951) About some interesting variation in the genus Laboulbenia. Biologisch Jaarboek (Dodonaea) 18: 122–129.

Middelhoek A. (1957) Eine neue Gattung der Laboulbeniales. Fungus 27: 72–75.

Miller MA, Pfeiffer W, Schwartz T. (2010) Creating the CIPRES science gateway for inference of large phylogenetic trees. Proceedings of the Gateway computing environments workshop (GCE), New Orleans, Louisiana 2010: 1–8. 10.1109/GCE.2010.5676129 DOI

Nguyen L-T, Schmidt HA, Von Haeseler A, Minh BQ. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. 10.1093/molbev/msu300 PubMed DOI PMC

Peyritsch J. (1873) Beiträge zur Kenntniss der Laboulbenien. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaft (Wien), Mathematisch-naturwissenschaftliche Classe 68: 227–254.

Pfliegler WP, Báthori F, Haelewaters D, Tartally A. (2016) Studies of Laboulbeniales on Myrmica ants (III): myrmecophilous arthropods as alternative hosts of Rickia wasmannii. Parasite 23: 1–50. 10.1051/parasite/2016060 PubMed DOI PMC

Rossi W. (2016) Contribution to the knowledge of the Laboulbeniales (Ascomycota) from Turkey. Webbia 71: 265–276. 10.1080/00837792.2016.1222668 DOI

Rossi W, Guéorguiev B, Georgiev G, Stoianova D. (2018) Laboulbeniales (Ascomycota) from Bulgaria and other countries. Plant Biosystems 153: 48–59. 10.1080/11263504.2018.1454531 DOI

Rossi W, Santamaría S. (2006) Laboulbenia casnoniae (Ascomycota, Laboulbeniales) and allied species. Nova Hedwigia 82: 189–204. 10.1127/0029-5035/2006/0082-0189 DOI

Rossi W, Weir A. (1997) Laboulbenia pterostichi and its allies. Mycological Research 101: 129–134. 10.1017/S0953756296002043 DOI

Sánchez-García M, Henkel TW, Aime MC, Smith ME, Matheny PB. (2016) Guyanagarika, a new ectomycorrhizal genus of Agaricales from the Neotropics. Fungal Biology 120: 1540–1553. 10.1016/j.funbio.2016.08.005 PubMed DOI

Santamaría S. (1998) Laboulbeniales, I. Laboulbenia. Flora Mycologica Iberica 4: 1–186.

Santamaría S. (2001) New and interesting Laboulbeniales (Fungi, Ascomycota) from Spain, IV. Nova Hedwigia 72: 375–389.

Santamaría S. (2003) Laboulbeniales, II. Acompsomyces-Ilyomyces. Flora Mycologica Iberica 5: 1–344.

Santamaría S, Balazuc J, Tavares II. (1991) Distribution of the European Laboulbeniales (Fungi, Ascomycotina). An annotated list of species. Treballs de l’Institut Botanic de Barcelona 14: 1–123.

Santamaría S, Cuesta Segura AD, Guardia L. (2020) New and remarkable Laboulbeniales (Ascomycota) from Spain. Nova Hedwigia 110: 347–367. 10.1127/nova_hedwigia/2020/0577 DOI

Santamaría S, Rossi W. (1993) Stigmatomyces (Laboulbeniales, Ascomycotina) ibéricos. Anales del Jardin Botanico de Madrid 51: 33–40.

Scheloske H-W. (1969) Beiträge zur Biologie, Ökologie und Systematik der Laboulbeniales (Ascomycetes) unter besondere Berücksichtigung des Parasit-Wirt-Verhältnisses. Parasitologische Schriftenreihe 19: 1–176.

Schoch CL, Sung GH, López-Giráldez F, Townsend JP, Miadlikowska J, Hofstetter V, Robbertse B, Matheny PB, Kauff F, Wang Z, Gueidan C, Andrie RM, Trippe K, Ciufetti LM, Wynns A, Fraker E, Hodkinson BP, Bonito G, Groenewald JZ, Arzanlou M, de Hoog GS, Crous PW, Hewitt D, Pfister DH, Peterson K, Gryzenhout M, Wingfield MJ, Aptroot A, Suh SO, Blackwell M, Hillis DM, Griffith GW, Castlebury LA, Rossman AY, Lumbsch HT, Lücking R, Büdel B, Rauhut A, Diederich P, Ertz D, Geiser DM, Hosaka K, Inderbitzin P, Kohlmeyer J, Volkmann-Kohlmeyer B, Mostert L, O’Donnell K, Sipman H, Rogers JD, Shoemaker RA, Sugiyama J, Summerbell RC, Untereiner W, Johnston PR, Stenroos S, Zuccaro A, Dyer PS, Crittenden PD, Cole MS, Hansen K, Trappe JM, Yahr R, Lutzoni F, Spatafora JW. (2009) The Ascomycota tree of life: A phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits. Systematic Biology 58: 224–239. 10.1093/sysbio/syp020 PubMed DOI

Singh G, Dal Grande F, Divakar PK, Otte J, Leavitt SD, Szczepanska K, Crespo A, Rico VJ, Aptroot A, da Silva Cáceres ME, Lumbsch HT, Schmitt I. (2015) Coalescentbased species delimitation approach uncovers high cryptic diversity in the cosmopolitan lichen-forming fungal genus Protoparmelia (Lecanorales, Ascomycota). PloS ONE 10: e0124625. 10.1371/journal.pone.0124625 PubMed DOI PMC

Spegazzini C. (1914) Primo contributo all a conoscenza delle Laboulbeniali italiani. Redia 10: 21–75.

Spegazzini C. (1915) Laboulbeniali ritrovate nelle collezioni di alcuni musei italiani. Anales del Museo Nacional de Historia Natural de Buenos Aires 26: 451–511.

Stefani FOP, Jones RH, May TW. (2014) Concordance of seven gene genealogies compared to phenotypic data reveals multiple cryptic species in Australian dermocyboid Cortinarius (Agaricales). Molecular Phylogenetics and Evolution 71: 249–260. 10.1016/j.ympev.2013.10.019 PubMed DOI

Sundberg H, Ekman S, Kruys Å. (2018a) A crush on small fungi: An efficient and quick method for obtaining DNA from minute ascomycetes. Methods in Ecology and Evolution 9: 148–158. 10.1111/2041-210X.12850 DOI

Sundberg H, Kruys Å, Bergsten J, Ekman S. (2018b) Position specificity in the genus Coreomyces (Laboulbeniomycetes, Ascomycota). Fungal Systematics and Evolution 1: 217–228. 10.3114/fuse.2018.01.09 PubMed DOI PMC

Tavares II. (1985) Laboulbeniales (Fungi, Ascomycetes). Mycologia Memoir 9: 1–627.

Thaxter R. (1890) On some North American species of Laboulbeniaceae. Proceedings of the American Academy of Arts and Sciences 25: 5–14. 10.2307/20020424 DOI

Thaxter R. (1891) Supplementary note on North American Laboulbeniaceae. Proceedings of the American Academy of Arts and Sciences 25: 261–270. 10.2307/20020441 DOI

Thaxter R. (1892) Further additions to the North American species of Laboulbeniaceae. Proceedings of the American Academy of Arts and Sciences 27: 29–45. 10.2307/20020464 DOI

Thaxter R. (1896) Contribution towards a monograph of the Laboulbeniaceae. Memoirs of the American Academy of Arts and Sciences 12: 187–429.

Thaxter R. (1899) Preliminary diagnoses of new species of Laboulbeniaceae. I. Proceedings of the American Academy of Arts and Sciences 35: 153–209. 10.2307/25129915 DOI

Thaxter R. (1900) Preliminary diagnoses of new species of Laboulbeniaceae. II. Proceedings of the American Academy of Arts and Sciences 35: 407–450. 10.2307/25129954 DOI

Thaxter R. (1901) Preliminary diagnoses of new species of Laboulbeniaceae. III. Proceedings of the American Academy of Arts and Sciences 36: 397–414. 10.2307/20021044 DOI

Thaxter R. (1908) Contribution toward a monograph of the Laboulbeniaceae. Part II. Memoirs of the American Academy of Arts and Sciences 13: 217–469. 10.2307/25058090 DOI

Thaxter R. (1931) Contribution toward a monograph of the Laboulbeniaceae V. Memoirs of the American Academy of Arts and Sciences 16: 1–435. 10.2307/25058136 DOI

Thiers B.2020(continuously updated)Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanic Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/ [accessed 14 July 2020]

Tragust S, Tartally A, Espadaler X, Billen J. (2016) Histopathology of Laboulbeniales (Ascomycota: Laboulbeniales): ectoparasitic fungi on ants (Hymenoptera: Formicidae). Myrmecological News 23: 81–89.

van Zuijlen JWA, Peeters TMJ, van Wielink PS, van Eck APW, Bouvy EHM. (1996) BRAND-STOF. Een inventarisatie van de entomofauna van het natuurreservaat ‘De Brand’ in 1990. Insektenwerkgroep KNNV, Tilburg, 228 pp.

Vilgalys R, Hester M. (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246. 10.1128/JB.172.8.4238-4246.1990 PubMed DOI PMC

Villarreal S RV, Valdés R LA, Bernal V JA, Piepenbring M, Kirschner R. (2010) A new species of Laboulbenia and new records from Panama. Mycological Progress 9: 57–67. 10.1007/s11557-009-0619-z DOI

Vorst O. (2010) Catalogue of the Coleoptera of the Netherlands. Monographieën van de Nederlandse Entomologische Vereniging 11: 5–317.

Walker MJ, Dorrestein A, Camacho JJ, Hiller T, Meckler LA, Silas KA, Haelewaters D. (2018) A tripartite survey of hyperparasitic fungi associated with ectoparasitic flies on bats (Mammalia: Chiroptera) in a neotropical cloud forest in Panama. Parasite 25: 1–19. 10.1051/parasite/2018017 PubMed DOI PMC

Weir A, Blackwell M. (2001b) Molecular data support the Laboulbeniales as a separate class of Ascomycota, Laboulbeniomycetes. Mycological Research 105: 715–722. 10.1016/S0953-7562(08)61989-9 DOI

Weir A, Hammond PM. (1997) Laboulbeniales on beetles: host utilization patterns and species richness of the parasites. Biodiversity and Conservation 6: 701–719. 10.1023/A:1018318320019 DOI

Weir A, Rossi W. (1995) Laboulbeniales parasitic on British Diptera. Mycological Research 99: 841–849. 10.1016/S0953-7562(09)80739-9 DOI

White TJ, Bruns TD, Lee SB, Taylor JW. (1990) Analysis of phylogenetic relationships by amplification and direct sequencing of ribosomal RNA genes. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ. (Eds) PCR Protocols: a Guide to Methods and Applications.Academic Press, San Diego, 315–322. 10.1016/B978-0-12-372180-8.50042-1 DOI

Wijayawardene NN, Hyde KD, Al-Ani LKT, Tedersoo L, Haelewaters D, Rajeshkumar KC, Zhao RL, Aptroot A, Leontyev DV, Saxena RK, Tokarev YS, Dai DQ, Letcher PM, Stephenson SL, Ertz D, Lumbsch HT, Kukwa M, Issi IV, Madrid H, Phillips AJL, Selbmann L, Pfliegler WP, Horváth E, Bensch K, Kirk PM, Kolaříková K, Raja HA, Radek R, Papp V, Dima B, Ma J, Malosso E, Takamatsu S, Rambold G, Gannibal PB, Triebel D, Gautam AK, Avasthi S, Suetrong S, Timdal E, Fryar SC, Delgado G, Réblová M, Doilom M, Dolatabadi S, Pawłowska J, Humber RA, Kodsueb R, Sánchez-Castro I, Goto BT, Silva DKA, de Souza FA, Oehl F, da Silva GA, Silva IR, Błaszkowski J, Jobim K, Maia LC, Barbosa FR, Fiuza PO, Divakar PK, Shenoy BD, Castañeda-Ruiz RF, Somrithipol S, Lateef AA, Karunarathna SC, Tibpromma S, Mortimer PE, Wanasinghe DN, Phookamsak R, Xu J, Wang Y, Tian F, Alvarado P, Li DW, Kušan I, Matočec N, Maharachchikumbura SSN, Papizadeh M, Heredia G, Wartchow F, Bakhshi M, Boehm E, Youssef N, Hustad VP, Lawrey JD, Santiago ALCMA, Bezerra JDP, Souza-Motta CM, Firmino AL, Tian Q, Houbraken J, Hongsanan S, Tanaka K, Dissanayake AJ, Monteiro JS, Grossart HP, Suija A, Weerakoon G, Etayo J, Tsurykau A, Vázquez V, Mungai P, Damm U, Li QR, Zhang H, Boonmee S, Lu YZ, Becerra AG, Kendrick B, Brearley FQ, Motiejūnaitė J, Sharma B, Khare R, Gaikwad S, Wijesundara DSA, Tang LZ, He MQ, Flakus A, Rodriguez-Flakus P, Zhurbenko MP, McKenzie EHC, Stadler M, Bhat DJ, Liu JK, Raza M, Jeewon R, Nassonova ES, Prieto M, Jayalal RGU, Erdoğdu M, Yurkov A, Schnittler M, Shchepin ON, Novozhilov YK, Silva-Filho AGS, Liu P, Cavender JC, Kang Y, Mohammad S, Zhang LF, Xu RF, Li YM, Dayarathne MC, Ekanayaka AH, Wen TC, Deng CY, Pereira OL, Navathe S, Hawksworth DL, Fan XL, Dissanayake LS, Kuhnert E, Grossart HP, Thines M. (2020) Outline of Fungi and fungus-like taxa. Mycosphere 11: 1060–1456. 10.5943/mycosphere/11/1/8 DOI

Yule GU. (1925) A mathematical theory of evolution based on the conclusions of Dr. J. C. Willis. Philosophical Transactions of the Royal Society B 213: 21–87. 10.1098/rstb.1925.0002 DOI

Zaneveld JS. (1938) Laboulbenia flagellata Peyritsch, een merkwaardige en voor Nederland nieuwe fungus. Fungus 9: 61–65.

New insights into the DNA extraction and PCR amplification of minute ascomycetes in the genus Laboulbenia (Pezizomycotina, Laboulbeniales)

The haustorium as a driving force for speciation in thallus-forming Laboulbeniomycetes

Hesperomyces (Fungi, Ascomycota) associated with Hyperaspis ladybirds (Coleoptera, Coccinellidae): Rethinking host specificity

Fungal Planet description sheets: 1284-1382