2026-03-032026-03-032025AMORIM, Dalton de Souza et al. An integrative taxonomic treatment of the Mycetophilidae (Diptera: Bibionomorpha) from Singapore reveals 115 new species on 730km2. Integrative Systematics, Stuttgartv, v. 8, n. 3, p. 1-318, 2025. DOI: 10.1101/2023.09.02.555672. Disponível em: https://bioone.org/journals/integrative-systematics-stuttgart-contributions-to-natural-history/volume-8/issue-3/2025.752376/An-integrative-taxonomic-treatment-of-the-Mycetophilidae-of-Singapore-reveals/10.18476/2025.752376.full. Acesso em: 2 mar. 2026.2692-8205https://bioone.org/journals/integrative-systematics-stuttgart-contributions-to-natural-history/volume-8/issue-3/2025.752376/An-integrative-taxonomic-treatment-of-the-Mycetophilidae-of-Singapore-reveals/10.18476/2025.752376.full“Open-ended” or “dark” taxa are species-rich clades so abundant and diverse that conventional taxonomic methods cannot keep pace with the volume of specimens and species. New approaches based on the presorting of specimens to putative species with low-cost DNA barcoding followed by morphological checks of a subset of specimens have made tackling these taxa more manageable. However, large scale integrative taxonomy will still require limiting the geographic scope of taxonomic revisions, given that most countries and biogeographic regions will have too many specimens and species for comprehensive coverage. Here, we present a case study of this type approach by carrying out a revision of the Mycetophilidae (Diptera), or fungus gnats, of Singapore with a reverse taxonomy workflow. The studied material was obtained from 496 samples collected with 71 Malaise traps placed at 107 sites in a variety of habitats in the country: mangroves, a swamp forest, freshwater swamp, primary rainforests, a coastal forest, and different types of secondary forest (old, maturing, young, urban). Based on molecular and morphological data for 1,454 specimens, we were able to delimit 120 species belonging to 23 genera. Of these, only five species had been previously described. The remaining 115 are new to science and described here. However, we only name 98 of these species, since 14 (Manota spp. A–G, Neoempheria spp. A–G) were represented solely by females from which we could not prepare a fully satisfactory morphological diagnosis, and three lacked molecular data (Epicypta sp. A, Epicypta sp. B, and Neoempheria sp. H). To assess congruence among species delimited by DNA barcodes (3% clusters) and morphology, we calculated match ratio and found it to be overall high across the entire family (95%), with even higher ratios (99%) for MOTUs clustered at 5% with Objective Clustering and MOTUs obtained with ABGD set to P=0.060. Overall, the ratio of undescribed to described species was an astonishing 20:1. Our revision of the Singapore fauna alone, we were able to increase the number of described Oriental Mycetophilidae species by about 25%, underscoring the magnitude of the taxonomic impediment for fungus gnats. Most of the Singapore Mycetophilidae belong to three genera: Neoempheria Osten-Sacken (29 species), Epicypta Winnertz (29 species), and Manota Williston (14 species) but we also describe a new genus, Integricypta Amorim & Oliveira gen. n. (Mycetophilinae: Mycetophilini), the putative sister-group of Aspidionia Colless, based on three species. The new species formally named here are: Leptomorphus rafflesi Amorim & Oliveira sp. n.; Monoclona simhapura Amorim & Oliveira sp. n.; Azana demeijeri Amorim & Oliveira sp. n.; Azana leekongchiani Amorim & Oliveira sp. n. (Sciophilinae); Tetragoneura crawfurdi Amorim & Oliveira sp. n.; Tetragoneura chola Amorim & Oliveira sp. n.; Tetragoneura dayuan Amorim & Oliveira sp. n.; Tetragoneura farquhari Amorim & Oliveira sp. n.; Ectrepesthoneura johor Amorim & Oliveira sp. n. (Tetragoneurinae); Mohelia zubirsaidi Amorim & Oliveira sp. n.; Allactoneura tumasik Amorim & Oliveira sp. n.; Allactoneura limbosengi Amorim & Oliveira sp. n.; Manota banzu Amorim & Oliveira sp. n.; Manota tantocksengi Amorim & Oliveira sp. n.; Manota bukittimah Amorim & Oliveira sp. n.; Manota chiamassie Amorim & Oliveira sp. n.; Manota danmaxi Amorim & Oliveira sp. n.; Manota mahuan Amorim & Oliveira sp. n.; Manota temenggong Amorim & Oliveira sp. n.; Clastobasis sritribuana Amorim & Oliveira sp. n.; Clastobasis bugis Amorim & Oliveira sp. n.; Clastobasis oranglaut Amorim & Oliveira sp. n. (Leiinae); Parempheriella mait Amorim & Oliveira sp. n.; Parempheriella longyamen Amorim & Oliveira sp. n.; Parempheriella peranakan Amorim & Oliveira sp. n.; Mycomya sachmatich Amorim & Oliveira sp. n.; Neoempheria merlio Amorim & Oliveira sp. n.; Neoempheria sabana Amorim & Oliveira sp. n.; Neoempheria sangabo Amorim & Oliveira sp. n.; Neoempheria shicheng Amorim & Oliveira sp. n.; Neoempheria ujong Amorim & Oliveira sp. n.; Neoempheria subaraji Amorim & Oliveira sp. n.; Neoempheria kokoiyeeae Amorim & Oliveira sp. n.; Neoempheria mandai Amorim & Oliveira sp. n.; Neoempheria malacca Amorim & Oliveira sp. n.; Neoempheria sinkapho Amorim & Oliveira sp. n.; Neoempheria singapura Amorim & Oliveira sp. n.; Neoempheria xinjiapo Amorim & Oliveira sp. n.; Neoempheria puluochung Amorim & Oliveira sp. n.; Neoempheria merdeka Amorim & Oliveira sp. n.; Neoempheria neesoon Amorim & Oliveira sp. n.; Neoempheria pulau Amorim & Oliveira sp. n.; Neoempheria cinkappur Amorim & Oliveira sp. n.; Neoempheria temasek Amorim & Oliveira sp. n.; Neoempheria polunini Amorim & Oliveira sp. n.; Neoempheria fajar Amorim & Oliveira sp. n.; Neoempheria riatanae Amorim & Oliveira sp. n. (Mycomyinae); Brachycampta glorialimae Amorim & Oliveira sp. n.; Brachycampta murphyi Amorim & Oliveira sp. n.; Brachycampta limtzepengi Amorim & Oliveira sp. n.; Rymosia teopohlengi Amorim & Oliveira sp. n.; Exechia tanswiehiani Amorim & Oliveira sp. n.; Exechia alinewongae Amorim & Oliveira sp. n.; Mycetophila chngseoktinae Amorim & Oliveira sp. n.; Mycetophila georgettechenae Amorim & Oliveira sp. n.; Mycetophila aishae Amorim & Oliveira sp. n.; Platyprosthiogyne phanwaithongae Amorim & Oliveira sp. n.; Platyprosthiogyne gohsookhimae Amorim & Oliveira sp. n.; Platyprosthiogyne rahimahae Amorim & Oliveira sp. n.; Platyprosthiogyne lynetteseahae Amorim & Oliveira sp. n.; Platyprosthiogyne neilaae Amorim & Oliveira sp. n.; Platyprosthiogyne snehalethaae Amorim & Oliveira sp. n.; Platurocypta adeleneweeae Amorim & Oliveira sp. n.; Platurocypta tanhoweliangi Amorim & Oliveira sp. n.; Epicypta constancesingamae Amorim & Oliveira sp. n.; Epicypta jennylauae Amorim & Oliveira sp. n.; Epicypta limchiumeiae Amorim & Oliveira sp. n.; Epicypta janetyeeae Amorim & Oliveira sp. n.; Epicypta kohkhenglianae Amorim & Oliveira sp. n.; Epicypta daintoni Amorim & Oliveira sp. n.; Epicypta holltumi Amorim & Oliveira sp. n.; Epicypta ridleyi Amorim & Oliveira sp. n.; Epicypta chezaharaae Amorim & Oliveira sp. n.; Epicypta tanjiakkimi Amorim & Oliveira sp. n.; Epicypta gehminae Amorim & Oliveira sp. n.; Epicypta jackieyingae Amorim & Oliveira sp. n.; Epicypta khatijunae Amorim & Oliveira sp. n.; Epicypta purchoni Amorim & Oliveira sp. n.; Epicypta foomaoshengi Amorim & Oliveira sp. n.; Epicypta ganengsengi Amorim & Oliveira sp. n.; Epicypta nanyangu Amorim & Oliveira sp. n.; Epicypta nus Amorim & Oliveira sp. n.; Epicypta peterngi Amorim & Oliveira sp. n.; Epicypta maggielimae Amorim & Oliveira sp. n.; Epicypta yupeigaoae Amorim & Oliveira sp. n.; Epicypta annwee Amorim & Oliveira sp. n.; Epicypta wallacei Amorim & Oliveira sp. n.; Epicypta lamtoongjini Amorim & Oliveira sp. n.; Epicypta catherinelimae Amorim & Oliveira sp. n.; Epicypta grootaerti Amorim & Oliveira sp. n.; Epicypta joaquimae Amorim & Oliveira sp. n.; Aspidionia cheesweeleeae Amorim & Oliveira sp. n.; Aspidionia janetjesudasonae Amorim & Oliveira sp. n.; Aspidionia fatimahae Amorim & Oliveira sp. n.; Integricypta fergusondavie Amorim & Oliveira sp. n.; Integricypta teosoonkimae Amorim & Oliveira sp. n.; Integricypta shirinae Amorim & Oliveira sp. n.; Integricypta hoyuenhoeae Amorim & Oliveira sp. n. (Mycetophilinae). The previously described species collected during this study are Metanepsia malaysiana Kallweit, Eumanota racola Søli, Parempheriella defectiva (Edwards), Neoempheria dizonalis (Edwards) (all known from Sumatra and/or the Malaysian peninsula), and Chalastonepsia hokkaidensis Kallweit, a species widespread in East Asia, whose female is described here for the first time. The mycomyine genus Vecella Wu & Yang is here proposed as a junior synonym to Parempheriella Matile, syn. n., with P. guadunana (Wu & Yang), comb. n. corresponding to an additional Palearctic species of Parempheriella. Barcodes for a second set of 1,493 Singapore mycetophilid specimens not analyzed here suggest the presence of an additional 18 MOTUs. We thus estimate that approximately 85% of all the Singapore species interceptable by Malaise traps are identified or described here. Our revision concludes with a discussion of the biogeography and generic composition of the Mycetophilidae fauna at the southern end of the Malay Peninsula. „Open-ended Taxa“oder „Dark Taxa“sind artenreiche Kladen, die so abundant und artenreich sind, dass die herkömmlichen taxonomischen Methoden angesichts der großen Anzahl von Exemplaren und Arten nicht greifen. Neue Ansätze, bei denen Exemplare zunächst mithilfe kostengünstiger DNA-Barcodes zu mutmaßlichen Arten vorsortiert und dann exemplarisch an einer kleinen Auswahl morphologisch nachgeprüft werden, haben die Bearbeitung dieser Taxa deutlich erleichtert. Allerdings kann auch mit DNA-Barcodes nur die Fauna eines vergleichsweise kleinen Gebietes bearbeiten werden, weil für die meisten Länder und biogeografischen Regionen zu viele Exemplare und Arten abgedeckt werden müssten. Wir demonstrieren hier in einer Fallstudie die Revision der Pilzmücken (Diptera: Mycetophilidae) von Singapur mittels eines umgekehrten taxonomischen Workflows. Das bearbeitete Material stammt aus 496 Proben, die mit 71 Malaise-Fallen an 107 Sammelstellen in verschiedenen Habitaten gefangen wurden: Mangroven, einem Sumpfwald, einem Süßwassersumpf, primäre Regenwälder, einem Küstenwald und verschiedene Sekundärwälder (jung, reifend, alt, urban). Basierend auf molekularen und morphologischen Daten von mehr als 1.454 Tieren grenzen wir 120 Arten aus 5 Gattungen ab, von denen lediglich 5 Arten bereits beschrieben sind. Die verbleibenden 115 Arten sind neu für die Wissenschaft und werden hier beschrieben. Allerdings benennen wir nur 98 Arten, da von 14 Arten (Manota spp. A–G, Neoempheria spp. A–G) derzeit nur Weibchen bekannt sind, für die derzeit keine zufriedenstellende morphologische Diagnose erstellt werden kann. Darüber hinaus fehlen molekulare Daten für drei Arten (Epicypta sp. A, Epicypta sp. B und Neoempheria sp. H). Was die Artgrenzen betrifft, so stimmen die molekularen und morphologischen Daten in den meisten Fällen überein (match ratio: 95% für 3% MOTUs). Eine noch höheres Übereinstimmungsverhältnis von 99% wird für 5% und ABGD MOTUs (P=0,060) beobachtet. Insgesamt ist das Verhältnis zwischen unbeschrieben und beschrieben erstaunlich hoch (20:1). Unsere Überarbeitung der Fauna Singapurs erhöht die Anzahl der beschriebenen Arten in der Orientalischen Region um über 25%, was das Ausmaß der taxonomischen Hürde für Pilzmücken unterstreicht. Die meisten der Mycetophilidenarten Singapurs gehören den drei Gattungen Neoempheria Osten-Sacken (29 Arten), Epicypta Winnertz (29 Arten) und Manota Williston (14 Arten), aber basierend auf drei Arten beschreiben wir hier auch eine neue Gattung, Integricypta Amorim & Oliveira gen. n., die zu den Mycetophilinae: Mycetophilini gehört und die mutmaßliche Schwestergruppe von Aspidionia Colless darstellt. Die sequenzierten, illustrierten hier formal benannten Arten sind Leptomorphus rafflesi Amorim & Oliveira sp. n.; Monoclona simhapura Amorim & Oliveira sp. n.; Azana demeijeri Amorim & Oliveira sp. n.; Azana leekongchiani Amorim & Oliveira sp. n. (Sciophilinae); Tetragoneura crawfurdi Amorim & Oliveira sp. n.; Tetragoneura chola Amorim & Oliveira sp. n.; Tetragoneura dayuan Amorim & Oliveira sp. n.; Tetragoneura farquhari Amorim & Oliveira sp. n.; Ectrepesthoneura johor Amorim & Oliveira sp. n. (Tetragoneurinae); Mohelia zubirsaidi Amorim & Oliveira sp. n.; Allactoneura tumasik Amorim & Oliveira sp. n.; Allactoneura limbosengi Amorim & Oliveira sp. n.; Manota banzu Amorim & Oliveira sp. n.; Manota tantocksengi Amorim & Oliveira sp. n.; Manota bukittimah Amorim & Oliveira sp. n.; Manota chiamassie Amorim & Oliveira sp. n.; Manota danmaxi Amorim & Oliveira sp. n.; Manota mahuan Amorim & Oliveira sp. n.; Manota temenggong Amorim & Oliveira sp. n.; Clastobasis sritribuana Amorim & Oliveira sp. n.; Clastobasis bugis Amorim & Oliveira sp. n.; Clastobasis oranglaut Amorim & Oliveira sp. n. (Leiinae); Parempheriella mait Amorim & Oliveira sp. n.; Parempheriella longyamen Amorim & Oliveira sp. n.; Parempheriella peranakan Amorim & Oliveira sp. n.; Mycomya sachmatich Amorim & Oliveira sp. n.; Neoempheria merlio Amorim & Oliveira sp. n.; Neoempheria sabana Amorim & Oliveira sp. n.; Neoempheria sangabo Amorim & Oliveira sp. n.; Neoempheria shicheng Amorim & Oliveira sp. n.; Neoempheria ujong Amorim & Oliveira sp. n.; Neoempheria subaraji Amorim & Oliveira sp. n.; Neoempheria kokoiyeeae Amorim & Oliveira sp. n.; Neoempheria mandai Amorim & Oliveira sp. n.; Neoempheria malacca Amorim & Oliveira sp. n.; Neoempheria sinkapho Amorim & Oliveira sp. n.; Neoempheria singapura Amorim & Oliveira sp. n.; Neoempheria xinjiapo Amorim & Oliveira sp. n.; Neoempheria puluochung Amorim & Oliveira sp. n.; Neoempheria merdeka Amorim & Oliveira sp. n.; Neoempheria neesoon Amorim & Oliveira sp. n.; Neoempheria pulau Amorim & Oliveira sp. n.; Neoempheria cinkappur Amorim & Oliveira sp. n.; Neoempheria temasek Amorim & Oliveira sp. n.; Neoempheria polunini Amorim & Oliveira sp. n.; Neoempheria fajar Amorim & Oliveira sp. n.; Neoempheria riatanae Amorim & Oliveira sp. n. (Mycomyinae); Brachycampta glorialimae Amorim & Oliveira sp. n.; Brachycampta murphyi Amorim & Oliveira sp. n.; Brachycampta limtzepengi Amorim & Oliveira sp. n.; Rymosia teopohlengi Amorim & Oliveira sp. n.; Exechia tanswiehiani Amorim & Oliveira sp. n.; Exechia alinewongae Amorim & Oliveira sp. n.; Mycetophila chngseoktinae Amorim & Oliveira sp. n.; Mycetophila georgettechenae Amorim & Oliveira sp. n.; Mycetophila aishae Amorim & Oliveira sp. n.; Platyprosthiogyne phanwaithongae Amorim & Oliveira sp. n.; Platyprosthiogyne gohsookhimae Amorim & Oliveira sp. n.; Platyprosthiogyne rahimahae Amorim & Oliveira sp. n.; Platyprosthiogyne lynetteseahae Amorim & Oliveira sp. n.; Platyprosthiogyne neilaae Amorim & Oliveira sp. n.; Platyprosthiogyne snehalethaae Amorim & Oliveira sp. n.; Platurocypta adeleneweeae Amorim & Oliveira sp. n.; Platurocypta tanhoweliangi Amorim & Oliveira sp. n.; Epicypta constancesingamae Amorim & Oliveira sp. n.; Epicypta jennylauae Amorim & Oliveira sp. n.; Epicypta limchiumeiae Amorim & Oliveira sp. n.; Epicypta janetyeeae Amorim & Oliveira sp. n.; Epicypta kohkhenglianae Amorim & Oliveira sp. n.; Epicypta daintoni Amorim & Oliveira sp. n.; Epicypta holltumi Amorim & Oliveira sp. n.; Epicypta ridleyi Amorim & Oliveira sp. n.; Epicypta chezaharaae Amorim & Oliveira sp. n.; Epicypta tanjiakkimi Amorim & Oliveira sp. n.; Epicypta gehminae Amorim & Oliveira sp. n.; Epicypta jackieyingae Amorim & Oliveira sp. n.; Epicypta khatijunae Amorim & Oliveira sp. n.; Epicypta purchoni Amorim & Oliveira sp. n.; Epicypta foomaoshengi Amorim & Oliveira sp. n.; Epicypta ganengsengi Amorim & Oliveira sp. n.; Epicypta nanyangu Amorim & Oliveira sp. n.; Epicypta nus Amorim & Oliveira sp. n.; Epicypta peterngi Amorim & Oliveira sp. n.; Epicypta maggielimae Amorim & Oliveira sp. n.; Epicypta yupeigaoae Amorim & Oliveira sp. n.; Epicypta annwee Amorim & Oliveira sp. n.; Epicypta wallacei Amorim & Oliveira sp. n.; Epicypta lamtoongjini Amorim & Oliveira sp. n.; Epicypta catherinelimae Amorim & Oliveira sp. n.; Epicypta grootaerti Amorim & Oliveira sp. n.; Epicypta joaquimae Amorim & Oliveira sp. n.; Aspidionia cheesweeleeae Amorim & Oliveira sp. n.; Aspidionia janetjesudasonae Amorim & Oliveira sp. n.; Aspidionia fatimahae Amorim & Oliveira sp. n.; Integricypta fergusondavie Amorim & Oliveira sp. n.; Integricypta teosoonkimae Amorim & Oliveira sp. n.; Integricypta shirinae Amorim & Oliveira sp. n.; Integricypta hoyuenhoeae Amorim & Oliveira sp. n. (Mycetophilinae). Die bereits beschriebenen Arten, die bei dieser Untersuchung gefangen wurden, sind Metanepsia malaysiana Kallweit, Eumanota racola Søli, Parempheriella defectiva (Edwards), Neoempheria dizonalis (Edwards) (alle derzeit aus Sumatra und/ oder der malaysischen Halbinsel bekannt), sowie Chalastonepsia hokkaidensis Kallweit, eine in Ostasien weit verbreitete Art, deren Weibchen hier erstmals beschrieben werden. Die Gattung Vecella Wu & Yang (Mycomyinae) wird hier als jüngeres Synonym für Parempheriella vorgeschlagen (syn. n.), wobei P. guadunana (Wu & Yang), comb. n. mit einer zusätzlichen palaearktischen Art von Parempheriella korrespondiert. Die hier nicht analysierten DNA-Barcodes einer zweiten Stichprobe aus 1.493 Exemplaren von Pilzmücken deuten darauf hin, dass in Singapur 18 zusätzliche MOTUs vorkommen. Daher schätzen wir, dass circa 85% aller Arten Singapurs, die regelmäßig in Malaise-Fallen geraten, hier beschrieben sind. Die Revision schließt mit einer Diskussion der Biogeographie und generischen Zusammensetzung der Mycetophilidenfauna im südlichen Teil der Malaiischen Halbinsel.engAcesso RestritoArtigo10.1101/2023.09.02.555672