Abstract
The current data on fossil empidoid flies of the subfamily Microphorinae (Dolichopodidae sensu lato) are summarized. The extinct taxa of this group includes five genera (Avenaphora Grimaldi & Cumming, Curvus Kaddumi, Meghyperiella Meunier, Microphorites Hennig, Pristinmicrophor Tang, Shi, Wang & Yang) and 17 described species (one species in extant genus Microphor Macquart and two species in Schistostoma Becker). One more possible fossil microphorine genus Microcathophorites is treated as a nomen nudum. Microphorines were found from different deposits dated over a very broad geological temporal range from Lower Cretaceous to the Eocene. However, the majority of them are currently known from the Cretaceous (five genera and 13 (14?) species). Only two genera and four (three?) species were found from the Eocene. More precise definitions of Curvus, Meghyperiella, Microphorites and Pristinmicrophor as well as their possible relationships are discussed. In addition, the little known Eocene genus Meghyperiella is redescribed and newly recorded from Rovno amber (Ukraine). An identification key to all fossil taxa of microphorines is compiled for the first time.
References
Alonso, J., Corral, J.C. & Martínez-Torres, L.M. (2005) Los yacimientos ambarígenos del Cretácico Inferior de Sierra Cantabria (Peñacerrada, Álava). In: Melendez, G. & Moreno-Azanza, M. (Eds.), La vida y los ambientes en el Periodo Cretácico. SEPAZ, pp. 81–142.
Arillo, A. (2007) Paleoethology: fossilized behaviours in amber. Geologica Acta, 5, 159–166.
Arillo, A., Peñalver, E. & Delclòs, X. (2008) Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits. Zootaxa, 1920 (1), 29–40. https://doi.org/10.11646/zootaxa.1920.1.2
Becker, T. (1902) Aegyptische Dipteren gesammelt und beschrieben. Mitteilungen aus dem Zoologischen Museum zu Berlin, II, 2, 1–66., pl. 1. https://doi.org/10.1002/mmnz.4830020237
Bramuzzo, S. & Nel, A. (2017) Youngest representative of the extinct genus Microphorites in the Eocene amber of France (Diptera: Dolichopodidae: Microphorinae). Zootaxa, 4231 (4), 590–594. https://doi.org/10.11646/zootaxa.4231.4.11
Brooks, S.E. & Cumming, J.M. (2022) Revision of the Nearctic Microphorinae (Diptera: Dolichopodidae sensu lato). Zootaxa, 5095 (1), 1–141. https://doi.org/10.11646/zootaxa.5095.1.1
Brooks, S.E., Cumming, J.M. & Grimaldi, D.A. (2019) Remarkable new fossil species of Schistostoma Becker (Diptera: Dolichopodidae: Microphorinae) from mid-Cretaceous Burmese amber. Zootaxa, 4624 (1), 121–131. https://doi.org/10.11646/zootaxa.4624.1.8
Chvála, M. (1983) The Empidoidea (Diptera) of Fennoscandia and Denmark. II. General Part. The families Hybotidae, Atelestidae and Microphoridae. Fauna Entomologica Scandinavica, 12, 1–279.
Chvála, M. (1986) Revision of Palaearctic Microphoridae (Diptera) 1. Microphor Macq. Acta Entomologica Bohemoslovaca, 83, 432–454.
Chvála, M. (1987) Revision of Palearctic Microphoridae (Diptera) 2. Schistostoma Beck. Acta Entomologica Bohemoslovaca, 84, 133–155.
Chvála, M. (1991) A new species of the Holarctic genus Schistostoma Becker (Diptera: Microphoridae) from South Africa, and first record of the family from South America. Annals of the Natal Museum, 32, 19–26.
Collin, J.E. (1960) Some Empididae from Palestine. Annals and Magazine of Natural History, 13, 385–420. https://doi.org/10.1080/00222935908650872
Collin, J.E. (1961) Empididae. In: British Flies. Vol. 6. University Press, Cambridge, 782 pp.
Cumming, J.M. & Brooks, S.E. (2002) Electrophorella, a new genus of parathalassiine flies from Baltic amber, with a cladistic analysis of the Microphorinae + Dolichopodidae lineage (Diptera: Empidoidea). Studia dipterologica, 9 (1), 41–54.
Cumming, J.M. & Brooks, S.E. (2019) Phylogenetic analysis and preliminary classification of the Parathalassiinae (Diptera: Empidoidea: Dolichopodidae sensu lato). Zootaxa, 4648 (1), 111–129. https://doi.org/10.11646/zootaxa.4648.1.5
Cumming, J.M. & Wood, D.M. (2017) [Chapter] 3. Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.), Manual of Afrotropical Diptera. Vol. 1. Introductory chapters and keys to Diptera families. Suricata 4. South African National Biodiversity Institute, Pretoria, pp. 89–133.
Dietrich, C.H., Dmitriev, D.A. & Perkovsky, E.E. (2021) A new leafhopper (Hemiptera: Cicadellidae) from late Eocene Rovno amber described based on the adult and associated last-instar nymph from Perebrody. Palaeoentomology, 4 (5), 445–452. https://doi.org/10.11646/palaeoentomology.4.5.10
Evenhuis, N.L. (1994) Catalogue of the Fossil Flies of the World (Insecta: Diptera). Backhuys Publishers, Leiden, 660 pp.
Fedotova, Z.A. & Perkovsky, E.E. (2015) New gall midges (Diptera, Cecidomyiidae, Stomatosematidi, Brachineuridi) from the Late Eocene amber of Gulyanka (Zhitomir Region, Ukraine). Paleontological Journal, 49, 270–278. https://doi.org/10.1134/S0031030115030041
Fedotova, Z.A. & Perkovsky, E.E. (2017) New genus and species of gall midges (Diptera, Cecidomyiidae, Porricondylinae, Holoneurini) from the Late Eocene amber of Olevsk (Zhitomir Region, Ukraine). Vestnik Zoologii, 51 (1), 23–30. https://doi.org/10.1515/vzoo-2017-0004
Giłka, W., Harbach, R.E. & Perkovsky, E.E. (2021) Mosquitoes (Diptera: Culicidae) in Eocene amber from the Rovno region, Ukraine. Zootaxa, 5016 (2), 257–270. https://doi.org/10.11646/zootaxa.5016.2.6
Grimaldi, D.A. & Cumming, J.M. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Ere-moneura. Bulletin of the American Museum of Natural History, 239, 1–124.
Grimaldi, D.A. & Engel, M.S. (2005) Evolution of the Insects. Cambridge University Press, New York, New York, 755 pp.
Hennig, W. (1971) Insektenfossilien aus der unteren Kreide. III. Empidiformia („Microphorinae“) aus der unteren Kreide und aus dem baltischen Bernstein; ein Vertreter der Cyclorrhapha aus der unteren Kreide. Stuttgarter Beiträge zur Naturkunde, 232, 1–28.
Jouault, C., Ngô-Muller, V., Zhang, Q. & Nel, A. (2020) New empidoid flies (Diptera: Atelestidae; Dolichopodidae) from mid-Cretaceous Burmese amber. Palaeoentomology, 3 (2), 204–211. https://doi.org/10.11646/palaeoentomology.3.2.10
Kaddumi, H.F. (2005) The Amber of Jordan. The Oldest Prehistoric Insects in Fossilized Resin. 2nd Edition. Eternal River Museum of Natural History, Amman, 224 pp.
Kaddumi, H.F. (2007) The Amber of Jordan. The Oldest Prehistoric Insects in Fossilized Resin. 3rd Edition. Eternal River Museum of Natural History, Amman, 298 pp.
Keilbach, R. (1982). Bibliographie und Liste der Arten tierischer Einschlüsse in fossilen Harzen sowie ihrer Aufbewahrungsorte. Deutsche entomologische Zeitschrift, Neue Folge 29, Heft 1–3 & 4–5, 129–286 & 301–391.
Kröber, O. (1912) Die Thereviden der indo-australischen region (Dipt.). Entomologische Mitteilungen, 1, 242–256.
Latreille, P.A. (1804) Nouveau dictionnaire d’histoire naturelle appliquée aux arts, à l’agriculture, à l’économie rurale et domestique, à la médecine, etc. par une société de naturalistes et d’agriculteurs. Tome vingt-quatrième. Tableau méthodiques d’histoire naturelle. Déterville, Paris, 238 pp.
Latreille, P.A. (1809) Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisique plurimis explicata. Tomus quartus et ultimus. A. Koenig, Parisiis et Argentorati, 399 pp. https://doi.org/10.5962/bhl.title.4819
Legalov, A.A., Nazarenko, V.Yu. & Perkovsky, E.E. (2021a) A new species of the genus Glaesotropis Gratshev & Zherikhin, 1995 (Coleoptera, Anthribidae) from Rovno amber. Fossil Record, 24, 1–7. https://doi.org/10.5194/fr-24-1-2021
Legalov, A.A, Nazarenko,V.Yu. & Perkovsky, E.E. (2021b) A new species of the genus Dorytomus Germar, 1817 (Coleoptera, Curculionidae) from Rovno amber. Zootaxa, 5006 (1), 95–100. https://doi.org/10.11646/zootaxa.5006.1.12
Legalov, A.A., Nazarenko,V.Yu., Vasilenko, D.V. & Perkovsky, E.E. (2022) Ceutorhynchus Germar (Coleoptera, Curculionidae) as proxy for Eocene core Brassicaceae: first record of the genus from Rovno amber. Journal of Paleontology, 96 (2), 379–386. https://doi.org/10.1017/jpa.2021.82
Linnaeus, C. (1758) Systema Naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis locis. Tomus I. Editio decima, reformata. Laurentius Salvius, Holmiæ [Stockholm], 824 pp. https://doi.org/10.5962/bhl.title.542
Lyubarsky, G. Yu. & Perkovsky, E.E. (2020) First Rovno amber species of the genus Telmatophilus (Coleoptera: Clavicornia: Cryptophagidae) from Veselukha floodplain, Invertebrate Zoology, 17 (1), 25–35. https://doi.org/10.15298/invertzool.17.1.03
Macquart, J. (1827) Insectes diptères du nord de la France. Platypézines, dolichopodes, empides, hybotides. Imprimerie de Leleux, Lille, 159 pp. https://doi.org/10.5962/bhl.title.148911
Macquart, J. (1834) Histoire naturelle des Insectes. Diptères. Tome premier. In: Roret, N.E. (Ed.), Collection des suites à Buffon, Paris, 1, pp. 1–578, 12 pls. https://doi.org/10.5962/bhl.title.14274
Martínez-Delclós, X., Briggs, D.E.G. & Peñalver, E. (2004) Taphonomy of insects in carbonates and amber. Palaeogeography, Palaeoclimatology, Palaeoecology, 203 (1–2), 19–64. https://doi.org/10.1016/S0031-0182(03)00643-6
Martínez-Torres, L.M., Alonso, J. & Valle, J.M. (2011) The Upper Aptian–Lower Albian amber deposit of the Peñacerrada II Geosite (Basque–Cantabrian Basin, Northern Spain): geological context and protection. Geoheritage, 3, 55–61. https://doi.org/10.1007/s12371-010-0030-9
Melander, A.L. (1928) Diptera, Fam. Empididae. In: Wytsman, P. (Ed.), Genera Insectorum, 185 (1927), pp. 1–434.
Melnitsky, S.I., Ivanov, V.D. & Perkovsky, E.E. (2021a) A new species of the fossil genus Electrotrichia (Insecta: Trichoptera: Hydroptilidae) from Rovno amber (Zhytomyr region, Olevsk locality). Palaeoentomology, 4 (5), 421–424. https://doi.org/10.11646/palaeoentomology.4.5.4
Melnitsky, S.I., Ivanov, V.D. & Perkovsky, E.E. (2021b) A new species of Plectrocnemia (Trichoptera: Polycentropodidae) from Rovno amber. Zootaxa, 5006 (1), 106–109. https://doi.org/10.11646/zootaxa.5006.1.14
Meunier, F. (1908) Monographie des Empidae de l’ambre de la Baltique et catalogue bibliographique complet sur les Diptères fossiles de cette résine. Annales des Sciences Naturelles, Zoologie, Series 9, 7, 81–135.
Negrobov, O.P. (1978) Flies of the superfamily Empidoidea (Diptera) from Cretaceous retinite in northern Siberia. Paleontologicheskii Zhurnal, 1978 (2), 81–90. [in Russian]
Nel, A., Garrouste, R. & Daugeron, C. (2017) Two new long-legged flies in the Santonian amber of France (Diptera: Dolichopodidae). Cretaceous Research, 69, 1–5. https://doi.org/10.1016/j.cretres.2016.08.009
Nel, A., Perrichot, V., Daugeron, C. & Néraudeau, D. (2004) A new Microphorites in the Lower Cretaceous amber of the Southwest of France (Diptera: Dolichopodidae, “Microphorinae”). Annales de la Société Entomologique de France, New Series, 40, 23–29. https://doi.org/10.1080/00379271.2004.10697401
Nentwig, W. (1985) Obligate kleptoparasitic behaviour of female flies at spider webs (Diptera: Empidoidea: Microphoridae). Zoologischer Anzeiger, 215, 348–354.
Ngô-Muller, V., Garrouste, R., Pouillon, J.-M. & Nel, A. (2020) A new species of the long-legged fly genus Microphorites in Burmese amber (Dolichopodidae: Microphorinae). Cretaceous Research, 107, 1–4. https://doi.org/10.1016/j.cretres.2019.104284
Perrichot, V. & Engel, M.S. (2014) Youngest occurrence of the genus Microphorites (Diptera: Dolichopodidae): a new species in Late Cretaceous Vendean amber. Paleontological Contributions, 10G, 30–33. https://doi.org/10.17161/PC.1808.15987
Radchenko, A.G., Perkovsky, E.E. & Vasilenko, D.V. (2021) Formica species (Hymenoptera, Formicidae, Formicinae) in late Eocene Rovno amber. Journal of Hymenoptera Research, 82, 237–251. https://doi.org/10.3897/jhr.82.64599
Rondani, C. (1856) Dipterologiae Italicae prodromus. Vol. 1. Genera Italica ordinis Dipterorum ordinatim disposita et distincta et in familias et stirpes aggregata. A. Storschi, Parmae [Parma], 228 + [2] pp. https://doi.org/10.5962/bhl.title.8160
Saigusa, T. & Yang, D. (2003) Empididae (Diptera) from Funiu Mountains, Henan, China (I). Studia dipterologica, 9 (2), 519–543. [2002]
Shamshev, I.V. & Sinclair, B.J. (2006) The genus Schistostoma Becker from southern Africa, with an evaluation of its generic status (Diptera: Dolichopodidae s.l.: Microphorinae). African Invertebrates, 47, 335–346.
Shamshev, I.V. (2020) First record of an extant species of the genus Schistostoma Becker (Diptera: Dolichopodidae: Microphorinae) from the Oriental Region. Russian Entomological Journal, 29 (3), 303–306. https://doi.org/10.15298/rusentj.29.3.11
Simutnik, S.A., Perkovsky, E.E., Khomych, M.R. & Vasilenko, D.V. (2021) First record of the Sulia glaesaria Simutnik, 2015 (Hymenoptera, Chalcidoidea, Encyrtidae) from Rovno amber. Journal of Hymenoptera Research, 88, 85–102. https://doi.org/10.3897/jhr.88.75941
Sinclair, B.J. & Cumming, J.M. (2006) The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). Zootaxa, 1180 (1), 1–172. https://doi.org/10.11646/zootaxa.1180.1.1
Solórzano Kraemer, M.M., Solórzano, K.A., Stebner, F., Bickel, D. & Rust, J. (2015) Entrapment bias of insects in Miocene amber revealed by trapping experiments in a tropical forest in Chiapas, Mexico. PlosOne, 10 (3), e.0118820. https://doi.org/10.1371/journal.pone.0118820
Solórzano Kraemer, M.M., Delclòs, X., Clapham, M., Arillo, A., Peris, D., Jäger, P., Stebner, F. & Peñalver, E. (2018) Arthropods in modern resins reveal if amber accurately recorded forest arthropod communities. Proceedings of the National Academy of Sciences, 115 (26), 6739–6744. https://doi.org/10.1073/pnas.1802138115
Spahr, U. (1985) Ergänzungen und Berichtigungen zu R. Keilbach’s Bibliographie und Liste der Bernsteinfossilien—Ordnung Diptera. Stutgarter Beiträge zur Naturkunde, Series B, 111, 1–146.
Tang, C.-f., Shi, C., Wang, S. & Yang, D. (2019) The first report of Dolichopodidae from mid-Cretaceous amber of northern Myanmar. Cretaceous Research, 104, 1–4. https://doi.org/10.1016/j.cretres.2019.07.009
Tkoč, M., Nel, A. & Prokop, J. (2016) Discovery of a new species of the Cretaceous genus Microphorites Hennig, 1971 (Diptera: Dolichopodidae s. lat.) in Paleogene amber from eastern Moravia (Czech Republic). Insect Systematics & Evolution, 47, 181–193. https://doi.org/10.1163/1876312X-47022139
Ulrich, H. (1991) Two new genera of parathalassiine-like flies from South Africa (Diptera: Empidoidea). Bonner zoologische Beiträge, 42, 187–216.
Ulrich, H. (2004) Phylogenetic considerations about an early colonization of the sea coasts by Dolichopodidae (Diptera). Studia dipterologica, 11, 233–243.
Yamamoto, S., Nazarenko, V.Y., Vasilenko, D.V. & Perkovsky, E.E. (2022) First fossil species of ship-timber beetles (Coleoptera: Lymexylidae) from Eocene Rovno amber (Ukraine). Fossil Record, 25 (1), 65–74. https://doi.org/10.3897/fr.25.81054
Zetterstedt, J.W. (1842) Diptera scandinaviae disposa et descripta. Vol. 1. ex Officina Lundbergiana, sumtibus auctoris, Lundae [Lund], 140 pp.