Abstract
Trachusa interrupta (Fabricius, 1781) s.l. has so far been regarded as a widespread resin bee in the tribe Anthidiini, whose range extends from northwest Africa and the Iberian Peninsula in the west to Central Asia and China in the east. It was thought that the pattern of yellow maculations on head, meso- and metasoma was quite variable. Several forms which were described since the 18th century have been regarded as synonyms. A comprehensive analysis of hundreds of specimens from all parts of its distribution found that T. interrupta s. l. actually represents a complex of closely related species. The study which included the examination of type material and the analyses of morphometric data of 15 measurements of the head, antennae and wings through multivariate statistical methods showed that there was relatively little variation in the colour pattern. Different colour patterns mostly represent different taxa which form distinctive clusters in Discriminant Function Analysis of morphometric data. The complex hereby consists of three widespread species, Trachusa interrupta (Fabricius, 1781) s.str., T. integra (Eversmann, 1852) stat. resurr., and T. anatolica sp. n., whose combined range extends from the Western Mediterranean to Central Asia and China, and five further species with restricted ranges in the southern part of the overall distribution: T. varia (Olivier, 1789) stat. resurr. and T. maghrebensis sp. n. in Spain and north-western Africa, and T. heinzi Dubitzky, 2007, T. grandicornis sp. n., and T. taurica sp. n. in Turkey and Iran. Additionally, some populations mainly of T. interrupta s. str. show in the southern part of its distribution (e.g. Spain, Italy, Greece) distinctive features in the colour pattern or morphological traits such as antennal length. As these characters are widely overlapping between populations and seem to follow geographic clines, these differences do not seem to reflect taxonomically relevant units. It was thought that they represent populations with reduced but still not ceased reproductive isolation and hence species in statu nascendi. All species of the T. interrupta complex as here defined have clearly delineated distributional areas. There is a little overlap in the distribution areas of the species and even the two most widespread species, T. interrupta s. str. and T. integra, which both occur widely in the West Palaearctic show mutually exclusive patchy distribution patterns, i.e. the two species normally do not occur in the same region. Nevertheless, a few cases were found where two species occur in sympatry, and a few cases (less than 1% of all specimens) where specimens had characters of two species. The latter may indicate that hybridisation occasionally occurs in parapatric contact zones.
References
Aguib, S. (2014) Biogéographie et Monographie des Megachilidae (Hymenoptera : Apoidea) dans le Nord Est Algérien. Thesis, Faculté des sciences de la Nature et de la Vie, Université Constantine 1, Constantine, 248 pp.
Aguib, S., Louadi, K. & Schwarz, M. (2010) Les Anthidiini (Megachilidae, Megachilinae) d’Algérie avec trois espèces nouvelles pour ce pays: Anthidium (Anthidium) florentinum (Fabricius, 1775), Anthidium (Proanthidium) amabile Alfken, 1932 et Pseudoanthidium (Exanthidium) enslini (Alfken, 1928). Entomofauna, 31, 121–152.
Amiet, F., Herrmann, M., Müller, A. & Neumeyer, R. (2004) Apidae 4: Anthidium, Chelostoma, Coelioxys, Dioxys, Heriades, Lithurgus, Megachile, Osmia, Stelis. In: Fauna Helvetica. Vol. 9. Centre Suisse de Cartographie de la Faune (CSCF) & Schweizerische Entomologische Gesellschaft (SEG), Neuchâtel, 274 pp.
Baldock, D., Wood, T., Cross, I. & Smit, J. (2018) The bees of Portugal (Hymenoptera: Apoidea: Anthophila). Entomofauna, 22 (Supplementum), 1–164.
Banaszak, J., Cierzniak, T., Kriger, R. & Wendzonka, J. (2006) Bees of xerothermic swards in the lower Vistula valley: diversity and zoogeographic analyses (Hymenoptera: Apoidea: Apiformes). Polish Journal of Entomology, 75, 105–154.
Costa, A (1884) Descrizione della specie nuove e nota sopra altre già conisciute. Rendiconti Accademia delle Scienze fisiche e matematiche Napoli, Serie 1a, 23, 13–31.
Dubitzky, A. (2007) Taxonomic notes on the western palaearctic species of Trachusa, subgenus Paraanthidium, with description of a new species from Turkey (Hymenoptera, Apoidea, Megachilidae). Mitteilungen der Münchener Entomologischen Gesellschaft, 97, 107–113.
Emlen, D. J. (2001) Costs and the diversification of exaggerated animal structures. Science, 291, 1534–1536.
https://doi.org/10.1126/science.1056607
Eversmann, E. (1852) Fauna Hymenopterologica Volgo-Uralensis. Bulletin de la Société Impériale des Naturalistes, Moscou, 25, 2–137.
Fabricius, J.C. (1787) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Tom 1. Christ. Gottl. Proft, Hafniae (København), 348 pp.
https://doi.org/10.5962/bhl.title.11657
Fabricius, J.C. (1793) Entomologia Systematica Emendata et aucta. Secundum Classes, Ordines, Genera, Species Adjectis Synonimis, Locis, Observationibus, Descriptionibus. Christ. Gottl. Proft, Hafniae (København), 519 pp.
https://doi.org/10.5962/bhl.title.122153
Fabricius, J.C. (1781) Species insectorum, exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosin, adjectis observationibus, descriptionibus. Tom 1. Hamburg & Kilonii, Bohnii, 552 pp.
https://doi.org/10.5962/bhl.title.36509
Fabricius, J.C. (1804) Systema Piezatorum Secundum Ordines, Genera, Species. Apud Carolum Reichard, Braunschweig (Brunsvigae), 439 + 30 pp.
Fateryga, A.V., Ivanov, S.P. & Filatov, M.A. (2018) Megachilid-bees (Hymenoptera: Megachilidae) of the Crimean Peninsula. Entomofauna, 39, 235–283.
Ferton, C. (1921) Notes détachées sur l’instinct des hyménoptères mellifères et ravisseurs avec la description de deux espèces nouvelles. Annales de la Société Entomologique de France, 89 (1920/1921), 329–375.
Friese, H. (1898) Die Bienen Europa‘s (Apidae Europaeae) nach ihren Gattungen, Arten und Varietäten auf vergleichend morphologisch-biologischer Grundlage. Theil IV. Solitäre Apiden: Genus Eriades. Genus Trachusa. Genus Anthidium. C. Lampe, Innsbruck, 303 pp.
Friese, H. (1911) Das Tierreich. Hymenoptera. Apidae I. Megachilinae. R. Friedländer und Sohn, Berlin, 440 pp.
Gmelin, J.F. & von Linné, C. (1790) Systema naturae per regna tria naturae : secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tom 1. Pars V. Georg Emanual Beer, Lipsiae (Leipzig), 27 pp. [pp. 2770–2796]
Grace, A. (2010) Introductory biogeography to bees of the Eastern Mediterranean and Near East. Bexhill Museum, Sussex, 284 pp.
Güler, Y., Dikmen, F., Töre, D. & Aytekin, A.M. (2014) Contributions on the current knowledge of the diversity of the Megachilidae (Apoidea: Hymenoptera) fauna in the Mediterranean Region of Turkey. Türkiye Entomoloji Dergisi, 38, 255–278.
https://doi.org/10.16970/ted.50880
Helbig, A.J., Knox, A.G., Parkin, D.T., Sangster, G. & Collinson, M. (2002) Guidelines for assigning species rank. Ibis, 144, 518–525.
https://doi.org/10.1046/j.1474-919X.2002.00091.x
Illiger, K. (1806) William Kirby‘s Familien der bienenartigen Insekten mit Zusätzen nach Weisungen und Bemerkungen. Magazin für Insektenkunde, 5, 28–175.
Kasparek, M. (2017a) Resin bees of the anthidiine genus Trachusa (Panzer, 1804): Identification, taxonomy, distribution and biology of the Old World species. Entomofauna, 21 (Supplementum), 1–152.
Kasparek, M. (2017b) The taxonomic identity of Anthidium fasciatellum Friese, 1917 (Hymenoptera: Apoidea: Anthidiini). Journal of Natural History, 51 (29–30), 1743–1757.
https://doi.org/10.1080/00222933.2017.1353152
Kasparek, M. (2018) Taxonomic revision proves Trachusa pubescens (Morawitz, 1872) sensu lato to be a complex of allopatric and sympatric species in South-Eastern Europe and Western Asia (Hymenoptera, Apoidea, Anthidiini). ZooKeys, 764, 111–144.
https://doi.org/10.3897/zookeys.764.24581
Kasparek, M. (2019) A new species in the Trachusa ovata species group (Hymenoptera: Apoidea) from Peninsular Malaysia with an overview of the Old World species within the genus Trachusa. Journal of Natural History. [53, 1079–1094]
https://doi.org/10.1080/00222933.2019.1632953
Latreille, P.A. (1809) Mémoire sur le genre l’Anthidie, Anthidium, de Fabricius classe des Insectes, ordre des Hyménoptères, famille des Apiaires. Annales du Muséum Histoire Naturelle, 13, 24–53.
Lepeletier, A. (1841) Histoire Naturelle des Insectes. Hyménoptères. Tom. 2. Saint-Germain-en-Laye, Paris, 680 pp.
Michener, C.D. (2007) The Bees of the World. Second edition. John Hopkins, Baltimore, 953 pp.
Müller, A. (1873) Über den Fundort von Anthidium curvipes Imhoff. Entomologische Zeitung (Stettinger Entomologische Zeitung), 34, 154–156.
Müller, A. (1996) Host-plant specialization in Western Palearctic Anthidine Bees (Hymenoptera: Apoidea: Megachilidae). Ecological Monographs, 66, 235–257.
https://doi.org/10.2307/2963476
Olivier, G.A. (1789) Abeille. Apis. Lin. Geof. Fab. [Linnaeus, Geoffroy, Fabricius]. Encyclopédie Méthodique ou par Ordre de Matières, Par une Société de Gens de Lettres, de Savans et d‘Artistes, Livr. Histoire Naturelle, Insectes, 4, 57–84.
Özbek, H. & van der Zanden, G. (1993) A preliminary review of the Megachilidae of Turkey. Part III. The Anthidiini (Hymenoptera: Apoidea). Türkiye Entomoloji Dergisi, 17, 193–207.
Praz, C. (2014) Zum aktuellen Zustand der Bienen. Bienen im Fokus von Wissenschaft und Politik. In: Bericht über das Symposium der Akademie der Naturwissenschaften Schweiz vom 26 November 2014. Swiss Academy of Sciences, Bern, pp. 9–10.
Proshchalykin, M.Yu., Astafurova, Yu.V. & Osytshnjuk, A.Z. (2017) The species-group names of bees (Hymenoptera: Apoidea, Apiformes) described from Crimea, North Caucasus, European part of Russia and Ural. Part II. Families Andrenidae and Megachilidae. Far Eastern Entomologist, 328, 1–34.
Samin, N. & Bağıraçık, N. (2016) A study on Crabronidae and Megachilidae (Hymenoptera: Apoidea) from West Azarbaijan province, Northwest of Iran. Entomofauna, 37, 493–504.
Saunders, E. (1908) Hymenoptera aculeata collected in Algeria by the Rev. A. E. Eaton, M. A., F. E. S., and the Rev. Francis David Morice, M. A., F. E. S. Transactions of the Entomological Society of London, 56, 177–274.
https://doi.org/10.1111/j.1365-2311.1908.tb02145.x
Schneider, D. (1964) Insect antennae. Annual Review of Entomology, 9, 103–122.
https://doi.org/10.1146/annurev.en.09.010164.000535
Schmid, W. (1872) Die schweizerischen Arten der Bienengattung Anthidium. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 3, 1869–1872, 448–475.
Schwarz, M., Gusenleitner, F., Westrich, P. & Dathe, H.H. (1996) Katalog der Bienen Österreichs, Deutschlands und der Schweiz (Hymenoptera, Apidae). Entomofauna, Supplementum, 8, 1–398.
Streinzer, M., Kelber, C., Pfabigan, S., Kleineidam, C.J. & Spaethe, J. (2013) Sexual dimorphism in the olfactory system of a solitary and a eusocial bee species. The Journal of Comparative Neurology, 521, 2742–2755.
https://doi.org/10.1002/cne.23312
van der Vecht, J. (1961) Hymenoptera Sphecoidea Fabriciana. Zoologische Verhandelingen, 48, 1–85.
Warncke, K. (1980) Die Bienengattung Anthidium Fabricius, 1804 in der Westpaläarktis und im turkestanischen Becken. Entomofauna, 1, 119–209.
Warncke, K. (1986) Die Wildbienen Mitteleuropas. Ihre gültigen Namen und ihre Verbreitung (Insecta: Hymenoptera). Entomofauna, 3 (Supplementum), 5–128.