Abstract
We have previously studied the musculature of the male genitalia in one species of Psilidae—Chyliza vittata Meigen, 1826. Here, we describe for the first time the male genital muscles of Imantimyia nigrifrons (Macquart, 1835), Loxocera aristata (Panzer, 1801), Chamaepsila humeralis (Zetterstedt, 1847), Psilosoma audouini (Zetterstedt, 1835), and Psila nigripalpis Shatalkin, 1983. It is shown that Psilidae are characterized by secondary symmetry of the pregenital and genital sclerites and muscles, and by reduction of the syntergosternite VII + VIII, which is correlated with reduction of the ejaculatory apodeme (apomorphy); the muscles M25 are also absent. The family shows compaction of the pregenital and genital skeleton and muscles realized in different ways. Clades [[[Chyliza] + [[Imantimyia + Loxocera] + [Chamaepsila + [Psilosoma + Psila]]] are supported by several synapomorphies.
Notes
Due to genital rotation, the sclerites of these segments do not always lie in the usual position; therefore, such characteristics as “wide” or “narrow” describe only the geometric shape of the sclerites, regardless of their orientation relative to the body axis.
REFERENCES
Barták, M. and Carles-Tolrá, M., Psilidae Loew, 1861, in Checklist of Diptera of the Czech Republic and Slovakia, Jedlička, L., Kúdela, M., and Stloukalová, V., Eds., 2009. Electronic version 2. http://www.edvis.sk/diptera2009/families/psilidae.htm
Brindle, A., Taxonomic notes on the larvae of British Diptera. 22 Psilidae, Entomologist, 1965, vol. 98, p. 169.
Buck, M., 60. Psilidae (rust flies, psilid flies), in Manual of Central American Diptera. Vol. 2, Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., and Zumbado, M.A., Eds., Ottawa: NRC Research Press, 2010, p. 837.
Buck, M. and Marshall, S., The identity of Pseudopsila, description of new subgenus of Psila, and redefinition of Psila sensu lato (Diptera: Psilidae), Eur. J. Entomol., 2006a, vol. 103, p. 183. https://doi.org/10.14411/eje.2006.021
Buck, M. and Marshall, S., Revision of New World Loxocera (Diptera: Psilidae), with phylogenetic redefinition of Holarctic subgenera and species groups, Eur. J. Entomol., 2006b, vol. 103, p. 193. https://doi.org/10.14411/eje.2006.022
Chandler, P.J., Observations on plant associations of the Psilidae, Entomol. Rec. J. Var., 1975, vol. 87, p. 13.
Chandler, P.J., Checklists of Insects of the British Isles (New Series) Part 1: Diptera, Handbooks for the Identification of British Insects. New Series. Vol. 12, London: Royal Entomological Society of London, 1998, p. 1.
Cogan, B.H., Family Psilidae, in A Catalog of the Diptera of the Oriental Region. Vol. 3, Delfinado, M.D. and Hardy, D.E., Eds., Hawaii: The University Press of Hawaii, Honolulu, 1977, p. 24.
Dengler, K., Zur Lebensweise von Chyliza leptogaster Panzer und Ch. nova Collin (Psilidae, Diptera) und ihre Bedeutung als Urheber von Bastnekrosen an Laubbäumen, Stud. Dipterol., 1997, vol. 4, p. 143.
Ellis, P.R., Hardman, J.A., and Saw, P.L., Host plants of the carrot fly, Psila rosae (F.) (Dipt., Psilidae), Entomol. Mon. Mag., 1992, vol. 128, p. 1.
Freidberg, A. and Shatalkin, A.I., The Psilidae (Diptera) of Israel, with emphasis on the Chamaepsila hebraica group, Isr. J. Entomol., 2009, vol. 38, p. 35.
Frey, R., Zur Systematik der paläarktischen Psiliden (Dipt.), Not. Entomol., 1925, vol. 5, p. 47.
Friedrich, F. and Beutel, R.G., The thorax of Zorotypus (Hexapoda, Zoraptera) and a new nomenclature for the musculature of Neoptera, Arthropod Struct. Dev., 2008, vol. 37, p. 29. https://doi.org/10.1016/j.asd.2007.04.003
Galinskaya, T.V. and Ovtshinnikova, O.G., Musculature of the male genitalia in the tribe Ulidiini (Diptera, Ulidiidae), Entomol. Rev., 2015a, vol. 95, no. 1, p. 31. https://doi.org/10.1134/S0013873815010042
Galinskaya, T.V. and Ovtshinnikova, O.G., Musculature of the male genitalia in Rivellia (Diptera: Platystomatidae), ZooKeys, 2015b, vol. 545, p. 149. https://doi.org/10.3897/zookeys.545.6702
Galinskaya, T.V., Shatalkin, A.I., and Ovtshinnikova, O.G., Skeleton and musculature of the male genitalia in the Nothybidae (Diptera), Entomol. Rev., 2016, vol. 96, no. 9, p. 1182. https://doi.org/10.1134/S001387381906006X
Galinskaya, T.V., Gafurova, D., and Ovtshinnikova, O.G., X-ray microtomography (microCT) of male genitalia of Nothybus kuznetsovorum (Nothybidae) and Cothornobata sp. (Micropezidae), ZooKeys, 2018, vol. 744, p. 139. https://doi.org/10.3897/zookeys.744.22347
Goloboff, P.A., Farris, J.S., and Nixon, K.C., TNT, a free program for phylogenetic analysis, Cladistics, 2008, vol. 24, p. 1. https://doi.org/10.1111/j.1096-0031.2008.00217.x
Goot, V.S. Van der and Veen, M.P. Van., De Spillebeenvliegen, Wortelvliegen en Wolzwevers van Noordwest-Europa, Utrecht, Nederland: Jeudbondsuitgeverij, 1996.
Hennig, W., Aussereuropäische Psiliden und Platystomiden im Deutschen Entomologischen Institut (Diptera), Arb. Morphol. Taxon. Entomol. Berlin-Dahlem, 1940, vol. 7, p. 304.
Hennig, W., 41. Psilidae, in Die Fliegen der Paläarktischen Region, Lindner, E., Ed., Stuttgart: Schweizerbart, 1941, p. 1.
Hennig, W., Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen, Beitr. Entomol., 1958, vol. 8, p. 505.
Iwasa, M., The Japanese species of the genus Chyliza (Diptera, Psilidae), Japan. J. Entomol., 1989, vol. 57, p. 148.
Iwasa, M., 3.11. Family Psilidae, in Contributions to a Manual of Palaearctic Diptera (with Special Reference to Flies of Economic Importance). Vol. 3. Higher Brachycera, Papp, L. and Darvas, B., Eds., Budapest: Science Herald, 1998, p. 177.
Lyneborg, L., On the life history of Chyliza annulipes Macquart, 1835 (Diptera: Psilidae), Entomol. Medd., 1987, vol. 55. p. 27.
Matsuda, R., Morphology and Evolution of the Insect Abdomen: with Special Reference to Developmental Patterns and Their Bearings upon Systematics, Oxford: Pergamon Press Ltd, 1976.
McAlpine, J.F., Morphology and terminology – Adults. 2, in Manual of Nearctic Diptera. Vol. 1, McAlpine, J.F., Ed., Ottawa: Research Branch, Agriculture Canada, 1981, p. 9.
McAlpine, J.F., Phylogeny and classification of the Muscomorpha, in Manual of Nearctic Diptera. Vol. 3, McAlpine, J.F., Ed., Ottawa: Research Branch, Agriculture Canada, 1989, p. 1397.
Meijere, J.C.H., de, Über die Larven der in Orchideen minierenden Dipteren, Tijdsch. Entomol., 1940, vol. 83, p. 122.
Meijere, J.C.H., de, Pupariën van Loxocera in stengels van Juncus, Entomol. Bericht., 1941, vol. 10, p. 286.
Nixon, K.C., ‘Winclada Version 1.00.08.’ 2002 (Published by the author, Ithaca, NY, USA). http://www.cladistics.com
Ovtshinnikova, O.G., Muscles of the male genitalia of Brachycera–Orthorrhapha (Diptera), Tr. Zool. Inst. Akad. Nauk SSSR, 1989, vol. 190, p. 1.
Ovtshinnikova, O.G., On the homology of male genital sclerites of Brachycera Orthorrhapha and Cyclorrhapha (Diptera) based on musculature, Dipterol. Res., 1994, vol. 5, no. 4, p. 263.
Ovtshinnikova, O.G. and Galinskaya, T.V., Musculature of the abdomen and male genitalia of a member of Celyphidae (Diptera, Cyclorrhapha), Oriental Insects, 2016a, vol. 50, no. 4, p. 178. http://dx.doi.org/10.1080/00305316.2016.1227733
Ovtshinnikova, O.G. and Galinskaya, T.V., Musculature of the male genitalia of a member of the genus Teleopsis Rondani, 1875 (Diopsidae, Diptera), Oriental Insects, 2016b, vol. 50, no. 2, p. 61. http://dx.doi.org/10.1080/00305316.2016.1171174
Ovtshinnikova, O.G., Galinskaya, T.V., and Shatalkin, A.I., Musculature of the male genitalia of Chyliza vittata Meigen (Diptera, Psilidae), Entomol. Rev., 2015, vol. 95, no. 7, p. 874. http://dx.doi.org/10.1134/S0013873815070052
Ovtshinnikova, O.G. and Yeates, D.K., Male genital musculature of Therevidae and Scenopinidae (Diptera: Asiloidea): structure, homology and phylogenetic implications, Aust. J. Entomol., 1998, vol. 37, no. 1, p. 27.
Pape, T., Fauna Europaea: Psilidae, in Fauna Europaea: Diptera, Brachycera, Fauna Europaea version 1.1, Pape, T., Ed., 2004. http://www.faunaeur.org
Shatalkin, A.I., Review of the East Palaearctic flies of the genus Psila Mg. (Diptera, Psilidae), with a key to the Palaearctic species, Tr. Zool. Inst. Akad. Nauk SSSR, 1986, vol. 146, p. 23.
Shatalkin, A.I., Notes on the Palaearctic Psilidae (Diptera), Archiv. Zool. Mus. Mosc. St. Univ., 1989, vol. 27, p. 88.
Shatalkin, A.I., Asian species of Loxocera Meigen (Diptera: Psilidae), Rus. Entomol. J., 1998, vol. 6, no. 3–4, p. 87.
Shatalkin, A.I., Afrotropical Psilidae (Diptera). 1. Genera Belobackenbardia gen. n. and Psila Meigen, 1803, Rus. Entomol. J., 2002, vol. 10, p. 417.
Shatalkin, A.I., 65. Psilidae (Psilid Flies, Naked Flies), in Manual of Afrotropical Diptera. Vol. 3. Brachycera: Cyclorrhapha, Excluding Calyptratae. Suricata 8, Kirk-Spriggs, A.H. and Sinclair, B.J., Eds., Pretoria: South African National Biodiversity Institute Publishing, 2021a, p. 1545.
Shatalkin, A.I., Belobackenbardia mostovskii sp. n. (Diptera: Psilidae) is the fourth species of an endemic genus found in South Africa, Rus. Entomol. J., 2021b, vol. 30, p. 342.
Shatalkin, A.I. and Merz, B., The Psilidae (Diptera, Acalyptrata) of Switzerland, with description of two new species from Central Europe, Rev. Suiss. Zool., 2010, vol. 117, p. 771. https://doi.org/10.5962/bhl.part.117602
Shewell, G.E., Family Psilidae, in A Catalog of the Diptera of America North of Mexico, Stone, A., Sabrosky, C.W., Wirth, W.W., Foote, R.H., and Coulson, J.R., Eds., Washington, D.C.: Smithsonian Press, 1965, p. 638.
Soós, A., Family Psilidae, in Catalogue of Palaearctic Diptera. Vol. 9, Soós, A. and Papp, L., Eds., Budapest: Akadémiai Kiadó, 1984, p. 28.
Steyskal, G.C., 60. Psilidae, in Manual of Nearctic Diptera. Vol. 2, McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R., and Wood, D.M., Eds., Ottawa: Research Branch, Agriculture Canada, 1987, p. 781.
Wiegmann, B.M., Trautwein, M.D., Winkler, I.S., Barr, N.W., Kim, J.-W., Lambkin, C., Bertone, M.A., Cassel, B.K., Bayless, K.M., Heimberg, A.M., Wheeler, B.M., Peterson, K.J., Pape, T., Sinclair, B.J., Skevington, J.H., Blagoderov, V., Caravas, J., Kutty, S., Schmidt-Ott, U., Kampmeier, G.E., Thompson, F.C., Grimaldi, D.A., Beckenbach, A.T., Courtney, G.W., Friedrich, M., Meier, R., and Yeates, D.K., Episodic radiations in the fly tree of life, Proc. Natl. Acad. Sci., 2011, vol. 108, no. 14, p. 5690. https://doi.org/10.1073/pnas.1012675108
Due to genital rotation, the sclerites of these segments do not always lie in the usual position; therefore, such characteristics as “wide” or “narrow” describe only the geometric shape of the sclerites, regardless of their orientation relative to the body axis.
ACKNOWLEDGMENTS
We are grateful to Lucia Pollini Paltrinieri (Museo cantonale di storia naturale, Lugano, Switzerland) for supplying the material, to D.I. Gavryushin and A.L. Ozerov (Zoological Museum, Lomonosov Moscow State University) for providing the material, the assistance and advice, useful criticism and many interesting discussions. We thank P.N. Petrov (Department of Entomology, Faculty of Biology, Lomonosov Moscow State University) for improving the English of the manuscript.
Funding
The work of A.I. Shatalkin was supported by the Scientific Project of the State order of the Government of the Russian Federation to Lomonosov Moscow State University no. 121032300105-0. The work of O.G. Ovtshinnikova was supported by the Zoological Institute of Russian Academy of Sciences (State Research Program no. 122031100272-3).
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Statement on the welfare of animals. All the applicable international, national, and/or institutional guidelines for the care and use of animals were followed. All the procedures performed in studies involving animals were in accordance with the ethical standards of the institution or practice at which the studies were conducted. The authors declare that they have no conflict of interest.
Additional information
Publisher's Note. Pleiades Publishing remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
About this article
Cite this article
Shatalkin, A.I., Ovtshinnikova, O.G. Skeleton and Musculature of the Male Genitalia in Psilidae (Diptera) and Their Implications for the Phylogeny. Entmol. Rev. 103, 373–400 (2023). https://doi.org/10.1134/S0013873823040012
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1134/S0013873823040012