This is the accepted version of the following article: Snežana Radenković, Ljiljana Šašić Zorić, Mihajla Djan, Dragana Obreht Vidaković, Jelena Ačanski, Gunilla Ståhls, Nevena Veličković, Zlata Markov, Theodora Petanidou, Nataša Kočiš Tubić, Ante Vujić (2018) Cryptic speciation in the Merodon luteomaculatus complex (Diptera: Syrphidae) from the eastern Mediterranean. Journal of Zoological Systematics and Evolutionary Research, 56(2): 170-191., which has been published in final form at [https://doi.org/10.1111/jzs.12193]. This article may be used for non-commercial purposes in accordance with the Wiley Self-Archiving Policy [https://authorservices.wiley.com/author-resources/Journal-Authors/licensing-open-access/open- access/self-archiving.html]. Journal of Zoological Systematics and Evolutionary Research Cryptic speciation in the Merodon luteomaculatus complex (Diptera: Syrphidae) from the Eastern Mediterranean Journal:For Journal Review of Zoological Systematics Only and Evolutionary Research Manuscript ID JZS.201700031.R3 Wiley - Manuscript type: Original Article Date Submitted by the Author: n/a Complete List of Authors: Radenkovic, Snezana; Faculty of Sciences, Department of Biology and Ecology Šašić Zorić, Ljiljana; University of Novi Sad, BioSense Institute Djan, Mihajla; Faculty of Sciences, Department of Biology and Ecology Obreht Vidaković, Dragana; University of British Columbia, Department of Forest and Conservation Sciences Ačanski, Jelena; University of Novi Sad, BioSense Institute Ståhls, Gunilla; Finnish Museum of Natural History, Zoology unit Veličković, Nevena; Faculty of Sciences, Department of Biology and Ecology Markov, Zlata; Faculty of Sciences, Department of Biology and Ecology Petanidou, Theodora; University of the Aegean, Department of Geography Kočiš Tubić, Nataša; Faculty of Sciences, Department of Biology and Ecology Vujić, Ante; Faculty of Sciences, Department of Biology and Ecology The Balkan Peninsula and Aegean region, hoverflies, DNA sequence data, Keywords: palaeogeography, wing and male genitalia geometric morphometry The Merodon aureus group is characterized by high endemism and the presence of morphologically cryptic species. Within one of its sub-groups, M. bessarabicus , seven species and four more species complexes have been described to date. One of these complexes, the M. luteomaculatus , comprises new taxa that are the subject of the present study. Its members have allopatric ranges restricted to the Balkan Peninsula and Aegean islands. This complex exhibits morphological variability that could not be characterized using a traditional morphological approach. Thus, we used Abstract: integrative taxonomy with independent character sets (molecular, geometric-morphometric, distributional) to delimit species boundaries. Data on three molecular markers ( COI , 28S rRNA , ISSR), geometric morphometry of the wing and male genitalia, together with distributional data, enabled recognition of six cryptic species within the complex: M. andriotes sp. n., M. euri sp. n., M. erymanthius sp. n., M. luteomaculatus sp. n., M. naxius sp. n. and M. peloponnesius sp. n. We discuss the possible influence of Aegean palaeogeographic history on the speciation of this complex. Page 1 of 84 Journal of Zoological Systematics and Evolutionary Research For Review Only Journal of Zoological Systematics and Evolutionary Research Page 2 of 84 1 Cryptic speciation in the Merodon luteomaculatus complex (Diptera: Syrphidae) from 2 the Eastern Mediterranean 3 Runing title: Merodon luteomaculatus complex 4 Snežana Radenković 1, Ljiljana Šašić Zorić 2,* , Mihajla Djan 1, Dragana Obreht Vidaković 3, 5 Jelena Ačanski 2, Gunilla Ståhls 4, Nevena Veličković 1, Zlata Markov 1, Theodora Petanidou 5, 6 Nataša Kočiš Tubić 1, Ante Vujić 1 7 1 Department of Biology and Ecology, Faculty of Sciences, University of Novi Sad, Trg 8 Dositeja ObradovićaFor 2, 21000 Review Novi Sad, Serbia. Only Phone: +381214852656, Fax: 9 +38121450620 10 2 BioSense Institute - Research Institute for Information Technologies in Biosystems, 11 University of Novi Sad, Dr Zorana Đinđića 1, 21000 Novi Sad, Serbia. Phone: +38121485 12 2137 13 3 Department of Forest and Conservation Sciences, University of British Columbia, #3027 – 14 2424 Main Mall, Vancouver, BC, V6T 1Z4. Phone: 604822 2507, Fax: 6048229102 15 4 Zoology Unit, Finnish Museum of Natural History, PO Box 17, 00014 University of 16 Helsinki, Finland. Phone: +3585031828824 17 5 Laboratory of Biogeography and Ecology, Department of Geography, University of the 18 Aegean, Mytilene, Greece. Phone: +302251036406, Fax: +30251036423. 19 20 *Corresponding author: [email protected] 21 22 23 Key words: The Balkan Peninsula and Aegean region, hoverflies, DNA sequence data, 24 palaeogeography, wing and male genitalia geometric morphometry. 1 Page 3 of 84 Journal of Zoological Systematics and Evolutionary Research 25 Abstract 26 The Merodon aureus group is characterized by high endemism and the presence of 27 morphologically cryptic species. Within one of its sub-groups, M. bessarabicus , seven 28 species and four more species complexes have been described to date. One of these 29 complexes, the M. luteomaculatus , comprises new taxa that are the subject of the present 30 study. Its members have allopatric ranges restricted to the Balkan Peninsula and Aegean 31 islands. This complex exhibits morphological variability that could not be characterized using 32 a traditional morphological approach. Thus, we used integrative taxonomy with independent 33 character sets (molecular,For geometric-morphometric,Review Only distributional) to delimit species 34 boundaries. Data on three molecular markers ( COI , 28S rRNA , ISSR), geometric 35 morphometry of the wing and male genitalia, together with distributional data, enabled 36 recognition of six cryptic species within the complex: M. andriotes sp. n., M. euri sp. n., M. 37 erymanthius sp. n., M. luteomaculatus sp. n., M. naxius sp. n. and M. peloponnesius sp. n. We 38 discuss the possible influence of Aegean palaeogeographic history on the speciation of this 39 complex. 2 Journal of Zoological Systematics and Evolutionary Research Page 4 of 84 40 Introduction 41 The hoverfly genus Merodon Meigen, 1803 comprises more than 160 species distributed over 42 Palaearctic and Ethiopian regions (Ståhls et al., 2009; Vujić et al., 2012). The most 43 comprehensive taxonomic study of this genus is that of Hurkmans (1993) who revised 61 44 Palaearctic species. During the last 15 years, multiple authors have significantly contributed 45 to clarifying the taxonomy and distribution of the taxa of this speciose genus, focussing on 46 particular species groups on the Iberian, Balkan and Anatolian Peninsulas (e.g. Marcos- 47 García, Vujić & Mengual, 2007; Ståhls et al., 2009; Popov, 2010; Radenković et al., 2011; 48 Vujić, Marcos-García,For Sarıbıyık Review & Ricarte, 2011; Vu Onlyjić et al., 2012; Vujić, Radenković, 49 Likov, Trifunov & Nikolić, 2013a; Popović et al., 2015; Vujić et al., 2015; Ačanski et al., 50 2016; Šašić et al., 2016; Veselić, Vujić & Radenković, 2017). It became the most species-rich 51 hoverfly genus in Europe (121 spp.) after the appearance of recent publications with newly 52 described species (Ačanski et al., 2016; Šašić et al., 2016; Veselić et al., 2017). This 53 phytophagous genus is especially widespread in the Mediterranean Basin, a region with a 54 high diversity of geophytes. Although the immature stages of only five species have been 55 described to date, and observational data on the host plants of larvae exist for only 15 species 56 (Andrić et al., 2014), it seems that the underground storage organs of geophytes are general 57 hosts for all Merodon species. 58 The taxonomy of one species group, named aureus , is especially challenging due to the 59 absence of distinct diagnostic characters like specific structure of genitalia or hind legs in 60 males. The shape of the male genitalia in this group is simple and very similar in all species 61 of the group. Its members can be easily recognized by their small size, rounded abdomen, and 62 the presence of a spike on the metatrochanter of males (Radenković et al., 2011; Šašić et al., 63 2016; Veselić et al., 2017). This group is characterized by high endemism (e.g. on the Iberian 64 Peninsula more than 70% of species from the aureus group are endemic to the region) and the 65 presence of morphologically cryptic species. Šašić et al. (2016) defined five sub-groups: M. 66 aureus Fabricius, 1805, M. bessarabicus Paramonov, 1924, M. chalybeus Wiedemann in 67 Meigen, 1822, M. cinereus (Fabricius, 1794) and M. dobrogensis Bradescu, 1982, in addition 68 to two species with independent positions, i.e. M. caerulescens Loew, 1869 and M. 69 unguicornis Strobl in Czerny & Strobl, 1909. Šašić et al. (2016) particularly focussed on 70 species delimitation within the atratus complex of the M. cinereus subgroup. Later, Veselić et 3 Page 5 of 84 Journal of Zoological Systematics and Evolutionary Research 71 al. (2017) reviewed the M. bessarabicus sub-group that is characterized by predominantly 72 pale tibiae and tarsi, and typically dark tergites. Authors distinguished seven species and 73 highlighted the presence of four additional species complexes the members of which could 74 not be differentiated by a traditional morphological taxonomic approach. One of these four 75 complexes is the M. luteomaculatus , which comprises new taxa that are the subject of the 76 present study. 77 Integrative taxonomy that