We provide the first molecular phylogeny of Keroplatidae and Lygistorrhinidae, families of fungus gnats (Diptera: Bibionomorpha: Sciaroidea). Phylogenies reconstructed by Maximum Likelihood and Bayesian methods, based on four nuclear and four mitochondrial gene markers (5106 base pairs) sequenced for 75 genera and 105 species, show Keroplatidae as monophyletic only with the family Lygistorrhinidae included, herewith treated as the subfamily Lygistorrhininae stat. nov. The subfamily Arachnocampinae is retained in the family, although lowering its overall support. An early branching clade, comprising species of Platyura Meigen, 1803 and Paleoplatyura melanderi Fisher, 1941, forms subfamily Platyurinae Loew, 1850 stat. nov. The subfamilies Sciarokeroplatinae and Macrocerinae grouped together with three genera considered here as Keroplatidae incertae sedis. Subfamily Lygistorrhininae forms a sister clade to subfamily Keroplatinae, both retained monophyletic with high support. The traditional division of the subfamily Keroplatinae into the tribes Orfeliini and Keroplatini appears as outdated, resting largely on adaptive characters prone to parallel evolution. We find support for an alternative tribe corresponding to the Cloeophoromyia-Asindulum genus group, but a tribal reclassification of the Keroplatinae is left for future studies. The genus Heteropterna Skuse, 1888 is considered as identical with Ctenoceridion Matile, 1972 syn. nov.

Department of Biology and Ecology Faculty of Science University of Ostrava Chittussiho 10 71000 Ostrava Czech Republic

Department of Life Sciences Natural History Museum Cromwell Road London SW7 5BD UK

Department of Natural Sciences National Museums Scotland Chambers Street Edinburgh EH1 1JF UK

Institute of Agricultural and Environmental Sciences Estonian University of Life Sciences Kreutzwaldi st 5D 51006 Tartu Estonia

Tromsø University Museum UiT The Arctic University of Norway P O box 6050 Langnes NO 9037 Tromsø Norway

Zobrazit více v PubMed

Ševčík J., Kaspřák D., Mantič M., Fitzgerald S., Ševčíková T., Tóthová A., Jaschhof M. Molecular phylogeny of the megadiverse insect infraorder Bibionomorpha sensu lato (Diptera) PeerJ. 2016;4:e2563. doi: 10.7717/peerj.2563. PubMed DOI PMC

Matile L. Recherches sur la systématique et lévolution des Keroplatidae: Diptera, Mycetophiloidea. Mem. Du Mus. Natl. D’histoire Nat. Ser. A Zool. 1990;148:1–682.

Evenhuis N. Catalog of The Keroplatidae of The World (Insecta: Diptera) Bishop Museum Press; Honolulu, HI, USA: 2006.

Ševčík J., Mantič M., Blagoderov V. Two new genera of Keroplatidae (Diptera), with an updated key to the World genera of Keroplatini. Acta Entomol. Musei Natl. Pragae. 2015;55:387–399.

Kurina O., Mantič M., Ševčík J. A Remarkable New Genus of Keroplatidae (Insecta, Diptera) from the Afrotropical Region, with DNA Sequence Data. Afr. Invertebr. 2017;58:93–105. doi: 10.3897/afrinvertebr.58.12655. DOI

Fungus Gnats Online. [(accessed on 22 April 2020)]; Available online: http://sciaroidea.info/

Evenhuis N. Catalogue of The Fossil Flies of the World (Insecta: Diptera) Backhuys; Leiden, The Netherlands: 1994.

Papp L., Ševčík J. Sciarokeroplatinae, a new subfamily of Keroplatidae (Diptera) Acta Zool. Acad. Sci. Hung. 2005;51:113–123. doi: 10.11646/zootaxa.2221.1.4. DOI

Blagoderov V., Grimaldi D. Fossil Sciaroidea (Diptera) In Cretaceous Ambers, Exclusive of Cecidomyiidae, Sciaridae, and Keroplatidae. Am. Mus. Novit. 2004;3433 doi: 10.1206/0003-0082(2004)433<0001:FSDICA>2.0.CO;2. DOI

Grimaldi D., Amorim D.D.S., Blagoderov V. The Mesozoic Family Archizelmiridae (Diptera: Insecta) J. Paleontol. 2003;77:368–381. doi: 10.1017/S0022336000043705. DOI

Baker C., Graham G., Scott K., Cameron S., Yeates D., Merritt D. Distribution and Phylogenetic Relationships of Australian Glow-Worms Arachnocampa (Diptera, Keroplatidae) Mol. Phylogenet. Evol. 2008;48:506–514. doi: 10.1016/j.ympev.2008.04.037. PubMed DOI

Saigusa T. Rare Sciaroid Genera from the Temperate east Palaearctic Region (Diptera: Sciaroidea) Jpn. J. Syst. Entomol. 2018;24:163–173.

Bechev D. Flower visitation of fungus gnats from the genera Antlemon, Asindulum and Macrorrhyncha (Diptera: Keroplatidae): Published data and a new record. ZooNotes. 2010;7:1–3.

Mantič M., Ševčík J. Macrocera rohaceki sp. nov. and other interesting records of Keroplatidae (Diptera) from southern and central Europe, with DNA sequence data. Acta Entomol. Musei Natl. Pragae. 2017;57:751–764. doi: 10.1515/aemnp-2017-0098. DOI

Reiter N., Freestone M., Brown G., Peakall R. Pollination by sexual deception of fungus gnats (Keroplatidae and Mycetophilidae) in two clades of Pterostylis (Orchidaceae) Bot. J. Linn. Soc. 2019;190:101–116. doi: 10.1093/botlinnean/boz009. DOI

Matile L. The Origin of Biodiversity in Insects: Phylogenetic Tests of Evolutionary Scenarios. 2nd ed. Volume 173. Memoires du Museum National d’Histoire Naturelle Seri, A Zoologie; Paris, France: 1997. Phylogeny and evolution of the larval diet in the Sciaroidea (Diptera, Bibionomorpha) since the Mesozoic; pp. 273–303.

Sharpe M.L., Dearden P.K., Gimenez G., Krause K.L. Comparative RNA seq analysis of the New Zealand glowworm Arachnocampa luminosa reveals bioluminescence-related genes. BMC Genom. 2015;16:825. doi: 10.1186/s12864-015-2006-2. PubMed DOI PMC

Falaschi R., Amaral D., Santos I., Domingos A., Johnson G., Martins A., Viroomal I., Pompéia S., Mirza J., Oliveira A., et al. Neoceroplatus betaryiensis nov. sp. (Diptera: Keroplatidae) is the first record of a bioluminescent fungus-gnat in South America. Sci. Rep. 2019;9 doi: 10.1038/s41598-019-47753-w. PubMed DOI PMC

Aiello A., Jolivet P. Myrmecophily in Keroplatidae (Diptera: Sciaroidea) J. N. Y. Entomol. Soc. 1996;104:226–230.

Chandler P.J., Matile L. A new species of Platyceridion Tollet (Diptera, Keroplatidae) with a larva predatory in ant infested internodes of Humboldtia laurifolia Vahl. Studia Dipterol. 1998;5:163–173.

Chandler P.J., Pijnakker J. Tropical fungus gnats established in nurseries in The Netherlands (Diptera: Keroplatidae) and Mycetophilidae. Br. J. Entomol. Nat. Hist. 2009;22:81.

Hickman V.V. On Planarivora insignis gen. et sp. n. (Diptera: Mycetophilidae), whose larval stages in land planarians are parasitic. Pap. Proc. R. Soc. Tasman. 1965;99:1–8.

Bertone M. Field Observations of Lygistorrhina sanctaecatharinae Thompson (Diptera: Sciaroidea) Proc. Entomol. Soc. Wash. 2018;120:448–451. doi: 10.4289/0013-8797.120.2.448. DOI

Edwards F. British Fungus-Gnats (Diptera, Mycetophilidae). With A Revised Generic Classification of the Family. Trans. R. Entomol. Soc. Lond. 1925;73:505–670. doi: 10.1111/j.1365-2311.1925.tb03369.x. DOI

Tuomikoski R. Systematic position of Lygistorrhina Skuse (Diptera, Mycetophiloidea) Ann. Entomol. Fenn. 1966;32:254–260.

Hennig W. Ordnung Diptera (Zweiflügler) Handb. Der Zool. 1973;4:1–337.

Søli G.E.E., Vokeroth J.R., Matile L. Families of Sciaroidea. In: Papp L., Darvas B., editors. Contributions to A Manual of Palaeartic Diptera. Science Herald; Budapest, Hungary: 2000. pp. 49–92.

Amorim D.D.S., Rindal E. Phylogeny of the Mycetophiliformia, with proposal of the subfamilies Heterotrichinae, Ohakuneinae, and Chiletrichinae for the Rangomaramidae (Diptera, Bibionomorpha) Zootaxa. 2007;1535:1–92. doi: 10.11646/zootaxa.1535.1.1. DOI

Hippa H., Vilkamaa P. Phylogeny of the Sciaroidea (Diptera): The Implication of additional taxa and character data. Zootaxa. 2006;1132:63–68. doi: 10.11646/zootaxa.1132.1.4. DOI

Wiegmann B.M., Trautwein M.D., Winkler I.S., Barr N.B., Kim J.-W., Lambkin C., Bertone M.A., Cassel B.K., Bayless K.M., Heimberg A.M., et al. Episodic radiations in the fly tree of life. Proc. Natl. Acad. Sci. USA. 2011;108:5690–5695. doi: 10.1073/pnas.1012675108. PubMed DOI PMC

Vilkamaa P., Hippa H. The genus Sciarotricha gen. n. (Sciaridae) and the phylogeny of recent and fossil Sciaroidea (Diptera) Insect Syst. Evol. 2005;36:121–143. doi: 10.1163/187631205788838492. DOI

Bertone M., Courtney G., Wiegmann B. Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes. Syst. Entomol. 2008;33:668–687. doi: 10.1111/j.1365-3113.2008.00437.x. DOI

Rindal E., Søli G.E., Bachmann L. Molecular phylogeny of the fungus gnat family Mycetophilidae (Diptera, Mycetophiliformia) Syst. Entomol. 2009;34:524–532. doi: 10.1111/j.1365-3113.2009.00474.x. DOI

Kallweit U. Lygistorrhina Skuse has branched from the Keroplatidae (Diptera: Sciaroidea)—A new perspective on the phylogenetic relationships in the keroplatid group of fungus gnats; Proceedings of the 8th International Congress of Dipterology; Potsdam, Germany. 10–15 August 2014; p. 156.

Thompson F.C. Notes on the genus Lygistorrhina Skuse with the description of the first Nearctic species (Diptera: Mycetophiloidea) Proc. Entomol. Soc. Wash. 1975;77:434–445.

Kaspřák D., Kerr P., Sýkora V., Tóthová A., Ševčík J. Molecular phylogeny of the fungus gnat subfamilies Gnoristinae and Mycomyinae, and their position within Mycetophilidae (Diptera) Syst. Entomol. 2019;44:128–138. doi: 10.1111/syen.12312. DOI

Sikora T., Jaschhof M., Mantič M., Kaspřák D., Ševčík J. Considerable congruence, enlightening conflict: Molecular analysis largely supports morphology-based hypotheses on Cecidomyiidae (Diptera) phylogeny. Zool. J. Linn. Soc. 2019;185:98–110. doi: 10.1093/zoolinnean/zly029. DOI

Katoh K., Standley D.M. MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usability. Mol. Biol. Evol. 2013;30:772–780. doi: 10.1093/molbev/mst010. PubMed DOI PMC

Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 2000;17:540–552. doi: 10.1093/oxfordjournals.molbev.a026334. PubMed DOI

Xia X., Lemey P. Assessing substitution saturation with DAMBE. Phylogenet. Handb. Pract. Approach DNA Protein Phylogeny. 2009;2:615–630.

Xia X. DAMBE5: A comprehensive software package for data analysis in molecular biology and evolution. Mol. Biol. Evol. 2013;30:1720–1728. doi: 10.1093/molbev/mst064. PubMed DOI PMC

Lanfear R., Frandsen P.B., Wright A.M., Senfeld T., Calcott B. PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Mol. Biol. Evol. 2016;34:772–773. doi: 10.1093/molbev/msw260. PubMed DOI

Yang Z. Among-site rate variation and its impact on phylogenetic analyses. Trends Ecol. Evol. 1996;11:367–372. doi: 10.1016/0169-5347(96)10041-0. PubMed DOI

Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., Von Haeseler A., Lanfear R. IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Mol. Biol. Evol. 2020;37:1530–1534. doi: 10.1093/molbev/msaa015. PubMed DOI PMC

Hoang D.T., Chernomor O., Haeseler A.V., Minh B.Q., Vinh L.S. UFBoot2: Improving the Ultrafast Bootstrap Approximation. Mol. Biol. Evol. 2017;35:518–522. doi: 10.1093/molbev/msx281. PubMed DOI PMC

Ronquist F., Huelsenbeck J.P. MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics. 2003;19:1572–1574. doi: 10.1093/bioinformatics/btg180. PubMed DOI

Miller M.A., Pfeiffer W., Schwartz T. Creating the CIPRES Science Gateway for inference of large phylogenetic trees; Proceedings of the 2010 gateway computing environments workshop (GCE); New Orleans, LA, USA. 14 November 2010; DOI

Rambaut A., Drummond A.J., Xie D., Baele G., Suchard M.A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 2018;67:901. doi: 10.1093/sysbio/syy032. PubMed DOI PMC

Letunic I., Bork P. Interactive Tree Of Life v2: Online annotation and display of phylogenetic trees made easy. Nucleic Acids Res. 2011;39:475–478. doi: 10.1093/nar/gkr201. PubMed DOI PMC

Edwards F.W. A note on the “New Zealand glow-worm” (Diptera, Mycetophilidae) Ann. Mag. Nat. Hist. 1924;14:175–179. doi: 10.1080/00222932408633104. DOI

Sabrosky C.W. Family-group names in Diptera: An annotated catalog. Myia. 1999;10:1–360.

Soós Á., Papp L. Catalogue of Palaearctic Diptera: Ceratopogonidae-Mycetophilidae. Elsevier; Amsterdam, The Netherlands: 1988.

Kurina O. Fungus Gnats in Estonia: (Diptera: Bolitophilidae, Keroplatidae, Macroceridae, Ditomyiidae, Diadocidiidae, Mycetophilidae) Tartu University Press; Tartu, Estonia: 1998.

Polevoi A.V. Fungus Gnats (Diptera: Bolitophilidae, Ditomyiidae, Keroplatidae, Diadocidiidae, Mycetophilidae) of Karelia. Karelian Research Centre, Russian Academy of Sciences; Petrozavodsk, Russia: 2000.

Papp L. New record of Diptera species from Hungary, with the list of the Hungarian Scathophagidae. Folia Entomol. Hung. 2006;67:121–128.

Papp L. Dixidae, Axymyiidae, Mycetobiidae, Keroplatidae, Macroceridae, and Ditomyiidae(Diptera) from Taiwan. Acta Zool. Acad. Sci. Hung. 2007;53:273–294.

Ševčík J. Langkawiana maculata gen. et sp. n. from Malaysia and its systematic position in the tribe Robsonomyiini (Diptera: Keroplatidae) Zootaxa. 2009;2221:58–66. doi: 10.11646/zootaxa.2221.1.4. DOI

Amorim D.D.S., Niu C., Li X., Lei C., Clarke A. Chetoneura shennonggongensis, a new species of cave-dwelling Keroplatini from China (Diptera: Keroplatidae), with a discussion of the position of Chetoneura. Zootaxa. 2008;1716:59–68. doi: 10.11646/zootaxa.1716.1.6. DOI

Blagoderov V. Lygistorrhinidae (Long-beaked Fungus Gnats) In: Kirk–Spriggs A.H., Sinclair B.J., editors. Manual of Afrotropical Diptera. Nematocerous Diptera and lower Brachycera. 2nd ed. Volume 2. South African National Biodiversity Institute; Pretoria, South Africa: 2017. pp. 527–531.

Hippa H., Mattsson I., Vilkamaa P. New Taxa of the Lygistorrhinidae (Diptera: Sciaroidea) and Their Implications for a Phylogenetic Analysis of the Family. Zootaxa. 2005;960:1–34. doi: 10.11646/zootaxa.960.1.1. DOI

Blagoderov V., Hippa H., Ševčík J. Asiorrhina, a new Oriental genus of Lygistorrhinidae (Diptera: Sciaroidea) and its phylogenetic position. Zootaxa. 2009;2295:31–45. doi: 10.11646/zootaxa.2295.1.4. DOI

Matile L. Revision des Truplaya Afrotropicaux (Diptera, Mycetophilidae) Ann. De La Société Entomol. De Fr. 1978;14:451–477.

Blagoderov V., Ševčík J. Keroplatidae (Predaceous Fungus Gnats) In: Kirk–Spriggs A.H., Sinclair B.J., editors. Manual of Afrotropical Diptera. Nematocerous Diptera and lower Brachycera. 2nd ed. Volume 2. South African National Biodiversity Institute; Pretoria, South Africa: 2017. pp. 527–531.

Matile L. Rocetelion, a new Holarctic genus of the Keroplatidae (Diptera, Mycetophiloidea): Description, phylogenetic and biogeographic notes. Ann. Entomol. Fenn. 1988;54:107–113.

Uesugi K. Fungus gnats of the genus Urytalpa Edwards (Diptera: Keroplatidae) in Japan. Entomol. Sci. 2004;7:369–376. doi: 10.1111/j.1479-8298.2004.00085.x. DOI

Kjærandsen J., Martinsson S., Hedmark K., Evenhuis N.L. On the genus Urytalpa Edwards (Diptera: Keroplatidae) in the Nordic and Nearctic Regions, with fixation of a new type species and a key to World males. Zootaxa. 2009;2160:29–50. doi: 10.11646/zootaxa.2160.1.2. DOI

Fitzgerald S. A Curious new genus of Keroplatini (Diptera: Keroplatidae) from Guatemala. Zootaxa. 2019;4586:383. doi: 10.11646/zootaxa.4586.2.11. PubMed DOI

Chandler P.J., Blasco-Zumeta J. The fungus gnats (Diptera, Bolitophilidae, Keroplatidae and Mycetophilidae) of the Monegros region (Zaragoza, Spain) and five other new European species of Pyratula Edwards and Sciophila Meigen. Zapateri. Rev. Aragonesa De Entomol. 2001;9:1–24. doi: 10.11646/zootaxa.2318.1.19. DOI

Kovac D., Matile L. Truplaya ferox (Insecta: Diptera: Mycetophiloidea), a new Malaysian keroplatid from bamboo phytotelmata with larvae predaceous on ants. Raffles Bull. Zool. 1997;45:15–28.

Matile L. Diptera: Mycetophilidae Keroplatinae. S. Afr. Anim. Life. 1974;15:1–532.

Ševčík J. Terocelion gen. nov., a new Oriental genus of Keroplatidae (Diptera) with pectinate antennae. Acta Entomol. Musei Natl. Pragae. 2012;52:495–503.

Davimacrocera, a New Extant Genus of Keroplatidae, with Notes on the Fossil Genus Burmacrocera (Diptera)

Extant Genus in the Mesozoic: Paleoplatyura Meunier (Diptera: Keroplatidae) Found in the Cretaceous Amber of Myanmar

Just a Fragment of Undescribed Diversity: Twenty New Oriental and Palearctic Species of Sciaroidea (Diptera), including DNA Sequence Data and Two New Fossil Genera

A Peculiar New Genus of Bibionomorpha (Diptera) with Brachycera-Like Modification of Antennae from Mid-Cretaceous Amber of Myanmar

Intriguing and Beautiful: Adamacrocera adami gen. et sp. nov. from the Upper Cretaceous Amber of Myanmar Represents a New Subfamily of Keroplatidae (Diptera: Bibionomorpha)