Review of the Merodon natans group with description of a new species, a key to the adults of known species of the natans lineage and first descriptions of some preimaginal stages Author Vujic, Ante https://orcid.org/0000-0002-8819-8079 University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovica 2, 21000 Novi Sad, Serbia; Ante Vujic [ante. vujic @ dbe. uns. ac. rs]; Tamara Tot [tamarat @ dbe. uns. ac. rs]; Sanja Veselic [sanja. veselic @ dbe. uns. ac. rs]; Maja Arok [maja. arok @ dbe. uns. ac. rs]; Snezana Radenkovic [snezana. radenkovic @ dbe. uns. ac. rs] Author Tot, Tamara https://orcid.org/0000-0001-8776-9362 University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovica 2, 21000 Novi Sad, Serbia; Ante Vujic [ante. vujic @ dbe. uns. ac. rs]; Tamara Tot [tamarat @ dbe. uns. ac. rs]; Sanja Veselic [sanja. veselic @ dbe. uns. ac. rs]; Maja Arok [maja. arok @ dbe. uns. ac. rs]; Snezana Radenkovic [snezana. radenkovic @ dbe. uns. ac. rs] Author Andric, Andrijana https://orcid.org/0000-0002-8239-7595 University of Novi Sad, BioSense Institute, Dr Zorana Đinđica 1, 21000 Novi Sad, Serbia; Andrijana Andric * [andrijana. andric @ biosens. uns. ac. rs]; Jelena Acanski [acanski @ biosense. rs]; Ljiljana Sasic Zoric [ljsasic @ biosense. rs] andrijana.andric@biosense.rs Author Acanski, Jelena University of Novi Sad, BioSense Institute, Dr Zorana Đinđica 1, 21000 Novi Sad, Serbia; Andrijana Andric * [andrijana. andric @ biosens. uns. ac. rs]; Jelena Acanski [acanski @ biosense. rs]; Ljiljana Sasic Zoric [ljsasic @ biosense. rs] Author Sasic Zoric, Ljiljana University of Novi Sad, BioSense Institute, Dr Zorana Đinđica 1, 21000 Novi Sad, Serbia; Andrijana Andric * [andrijana. andric @ biosens. uns. ac. rs]; Jelena Acanski [acanski @ biosense. rs]; Ljiljana Sasic Zoric [ljsasic @ biosense. rs] Author Perez-Banon, Celeste Department of Environmental Sciences and Natural Resources, Faculty of Sciences III, Campus of San Vicente, University of Alicante, Alicante, Spain; Celeste Perez-Banon [celeste. perez @ ua. es]; Andrea Aracil [and. aracilgisbert @ gmail. com]; Santos Rojo [santos. rojo @ ua. es] Author Aracil, Andrea Department of Environmental Sciences and Natural Resources, Faculty of Sciences III, Campus of San Vicente, University of Alicante, Alicante, Spain; Celeste Perez-Banon [celeste. perez @ ua. es]; Andrea Aracil [and. aracilgisbert @ gmail. com]; Santos Rojo [santos. rojo @ ua. es] Author Veselic, Sanja University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovica 2, 21000 Novi Sad, Serbia; Ante Vujic [ante. vujic @ dbe. uns. ac. rs]; Tamara Tot [tamarat @ dbe. uns. ac. rs]; Sanja Veselic [sanja. veselic @ dbe. uns. ac. rs]; Maja Arok [maja. arok @ dbe. uns. ac. rs]; Snezana Radenkovic [snezana. radenkovic @ dbe. uns. ac. rs] Author Arok, Maja University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovica 2, 21000 Novi Sad, Serbia; Ante Vujic [ante. vujic @ dbe. uns. ac. rs]; Tamara Tot [tamarat @ dbe. uns. ac. rs]; Sanja Veselic [sanja. veselic @ dbe. uns. ac. rs]; Maja Arok [maja. arok @ dbe. uns. ac. rs]; Snezana Radenkovic [snezana. radenkovic @ dbe. uns. ac. rs] Author Mengual, Ximo https://orcid.org/0000-0002-6185-9404 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D- 53113 Bonn, Germany; Ximo Mengual [x. mengual @ leibniz-zfmk. de] Author van Eck, Andre BioMongol Foundation, Korte Hoefstraat 30, 5046 DB Tilburg, The Netherlands; Andre van Eck [andrevaneck @ biomongol. org] Author Rojo, Santos https://orcid.org/0000-0003-2160-9643 Department of Environmental Sciences and Natural Resources, Faculty of Sciences III, Campus of San Vicente, University of Alicante, Alicante, Spain; Celeste Perez-Banon [celeste. perez @ ua. es]; Andrea Aracil [and. aracilgisbert @ gmail. com]; Santos Rojo [santos. rojo @ ua. es] Author Radenkovic, Snezana University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovica 2, 21000 Novi Sad, Serbia; Ante Vujic [ante. vujic @ dbe. uns. ac. rs]; Tamara Tot [tamarat @ dbe. uns. ac. rs]; Sanja Veselic [sanja. veselic @ dbe. uns. ac. rs]; Maja Arok [maja. arok @ dbe. uns. ac. rs]; Snezana Radenkovic [snezana. radenkovic @ dbe. uns. ac. rs] text Arthropod Systematics & amp; Phylogeny 2021 2021-08-10 79 343 378 http://dx.doi.org/10.3897/asp.79.e65861 journal article http://dx.doi.org/10.3897/asp.79.e65861 1864-8312-79-343 F45452139C354104A97E14A8D1AC8029 B9CFCA6CEFAD57339B463739D73914C7 Merodon makrisi Vujic , Radenkovic & Tot sp. nov. Fig. 7A, B Merodon aff. natans in van Steenis et al. 2019 : 139. Type locality. CYPRUS: Limassol, Platres, Trooditissa Picnic Site, 4 Oct. 2017, X. Mengual leg. Type material. Holotype : 1 ♂, pinned, with genitalia in a separate microvial with glycerine. Left metaleg glued to the locality label. Original labels: "Cyprus: Limassol, Platres, / Trooditissa Picnic Site, 1340m., / 34.914736°N 32.842261°E , / 4 Oct. 2017. X. Mengual leg.", "DNA voucher specimen / ZFMK, Lab code / D363 / Bonn, Germany", " ZFMK DIP / 00028063 [QR code]". Paratypes : CYPRUS: 1 ♂; Mia Milia; 2 Nov. 1926; S.J. Curry leg.; "Anti Locust Res. Centre Reg. No. 453"; FSUNS ID 04973 (TAUI) 1 ♀; same data as for preceding; FSUNS ID 04965 (TAUI) 1 ♀; Limassol, Eftagonya; 550 m a.s.l.; 25 Oct. 1951; G. Mavromoustakis leg.; KBIN (R.I.Sc.N.B. 24.236) 3 ♂♂; Paramali; 34.6754°N , 32.8043°E ; 1 Nov. 2013; C. Makris leg.; AvE coll. 1 ♀; Limassol, Fasouri; 34.63°N , 32.92°E ; 9 Nov. 2013; C. Makris leg.; AvE coll. 5 ♂♂; Episkopi; 34.87°N , 32.87°E ; 4 Nov. 2014; C. Makris leg.; AvE coll. 1 ♂; Episkopi, Kourion; 34.6699°N , 32.8754°E ; 7 Feb. 2016; A. van Eck leg.; collected bulbs of Prospero autumnale ( Scilla autumnalis ), 1 male emerged 21 Oct. 2016; CEUA 1 ♀; Episkopi; 34.8754°N , 32.8754°E ; 30 Oct. 2016; A. van Eck leg.; AvE coll. 2 ♂♂; Lefkara; 34.9026°N , 33.3392°E ; 31 Oct. 2016; A. van Eck leg.; on low hanging branches and leaves of carob tree; AvE coll. 1 ♀; Episkopi; 34.8754°N , 32.8754°E ; 2 Nov. 2016; A. van Eck leg.; on bare ground, in Pinus brutia / Cistus -vegetation; AvE coll. 1 ♂; same data as for preceding; on the ground, in more open vegetation 1 ♀; Neo Chorio, Akamas; 35.0259°N , 32.3508°E ; 200 m a.s.l.; 2 Nov. 2016; A. van Eck leg.; AvE coll. 1 ♂; Kakopetria; 34.9746°N , 32.9132°E ; 3 Nov. 2016; A. van Eck leg.; AvE coll. 2 ♂♂, 2 ♀♀; Agios Sozomenos; 35.0496°N , 33.4406°E ; 5 Nov. 2016; C. Makris leg.; AvE coll. 1 ♂, 3 ♀♀; Lemesos; 34.66°N , 32.87°E ; 4 Oct. 2017; C. Makris leg.; near Prospero autumnale ; AvE coll. 1 ♂; Limassol, Platres, Trooditissa Picnic Site; 34.9147°N , 32.8422°E ; 1340 m a.s.l.; 4 Oct. 2017; X. Mengual leg.; ZFMK-DIP-00028063 (DNA-voucher D363) (ZFMK) in van Steenis et al. (2019) as Merodon aff. natans ; 1 ♂; Episkopi, near Ancient Kourion stadium; 34.6709°N , 32.8745°E ; 112 m a.s.l.; 7 Oct. 2017; X. Mengual leg.; ZFMK-DIP-00028064 (DNA-voucher D367) (ZFMK) in van Steenis et al. (2019) as Merodon aff. natans ; 1 ♀; Paphos, Paphos Forest, Appides stream; 34.9913°N , 32.647°E ; 747 m a.s.l.; 6-12 Oct. 2017; X. Mengual leg.; double-head Malaise trap; ZFMK-DIP-00027887 (DNA-voucher D395) (ZFMK) 1 ♂; Kyrenia, Lapithos; 35.3477°N , 33.1504°E ; 5 m a.s.l.; 14-28 Oct. 2018; Oe . Oezden leg.; Malaise trap in garden; AvE coll.; ( Merodon sp. in van Eck et al. (2020b) ). - Additional material examined . ISRAEL: 2 ♂♂; Mount Carmel; 32.7430°N , 35.0475°E ; 30 Sep. 1971; A. Freidberg leg.; det. W. Hurkmans as Merodon natans ; TAUI 1 ♀; Tel Aviv country club; 32.0780°N , 34.8128°E ; 16 Nov. 1974; M. Yakubowski leg.; det. W. Hurkmans as Merodon natans ; TAUI 12 ♂♂, 7 ♀♀; Nahal Poleg; 32.2452°N , 34.8580°E ; 7 Nov. 1977; M. Kaplan leg.; det. W. Hurkmans as Merodon natans ; TAUI 4 ♂♂; same data as for preceding; FSUNS ID 04892, 04893, 04950, 04951 (TAUI) 4 ♂♂, 8 ♀♀; same data as for preceding; A. Freidberg leg; TAUI 1 ♀; same data as for preceding; FSUNS ID 04955 (TAUI) 6 ♂♂; Yarhiv; 32.1490°N , 34.9674°E ; 3 Nov. 1977; M. Kaplan leg.; det. W. Hurkmans as Merodon natans ; TAUI 1 ♂; same data as for preceding; FSUNS ID 04956 (TAUI) 3 ♀♀; same data as for preceding; FSUNS ID 04894, 04953, 04954 (TAUI) 1 ♂; same data as for preceding; A. Freidberg leg.; FSUNS ID 04952 (TAUI) 1 ♂; Tel Aviv; 32.0780°N , 34.8128°E ; 7 Nov. 1977; M. Kaplan leg.; det. W. Hurkmans as Merodon natans ; TAUI 1 ♂; Ben-Shemen; 31.9516°N , 34.9353°E ; 13 Nov. 1982; Ben-Shahar leg.; TAUI 1 ♂, 1 ♀; Jerusalem; 31.7784N, 35.2080E; 18 Oct. 1988; R. Kasher leg.; det. W. Hurkmans as Merodon natans ; TAUI 1 ♂; Ayalon Park Canada; 31.8362°N , 34.9955°E ; 4 Nov. 1996; L. Friedman leg.; TAUI 1 ♀; same data as for preceding; FSUNS ID 04895 (TAUI) 1 ♀; Yakum; 32.2508°N , 34.8383°E ; 19 Oct. 1996; Hofman leg.; TAUI. - SOMALIA: 1 ♀; Galgala Oasis; 11.000°N , 49.050°E ; 14 Oct. 1973; collected by the Spedizione Biologica in Somalia, 1973; MZUF 1 ♀; plateau between Hongolo and Bur Inaoski; 9.250°N , 50.330°E ; 1924; G. Stefanini, N. Puccioni leg.; MZUF. Figure 7. Merodon makrisi Vujic , Radenkovic & Tot sp. nov. A male in Cyprus, photo: C. Makris, 14 Nov. 2016 B female in Cyprus, photo: C. Makris, 24 Oct. 2013 C typical habitat in Cyprus where M. makrisi Vujic , Radenkovic & Tot sp. nov. can be collected. Bulbs of Prospero autumnale were collected in the patch in the front, from which one male specimen was reared. Photo: A. van Eck, 7 Feb. 2016. Diagnosis. Morphological features such as a large fossette (Fig. 2E ) and golden erect pile on terga in male (Fig. 7A ); position of fossette close to base of arista (Fig. 2I :f); and dense white pile on eyes in both sexes separate this species from other species of the Merodon natans group. Taxonomic notes. Merodon makrisi sp. nov. is a medium sized species (9-13 mm). Morphological features such as a longer basoflagellomere (Fig. 2E, F ); and broader metafemur (Fig. 4E, F ) in both sexes, and large, rounded posterior surstyle lobe (Figs 8A :p, B:p, 10D) of male genitalia, easily separate this species from the related M. calcaratus (in M. calcaratus posterior surstyle lobe narrow, fingerlike (Fig. 5A-G )). Merodon makrisi sp. nov. is very similar to both M. natans and M. pulveris from which males can be distinguished by a set of characters of antenna, shape of the male genitalia and pile on terga and eyes: fossette in M. makrisi sp. nov. large, extending from the base of arista to the apex of basoflagellomere (Fig. 2E ) (fossette in M. natans and M. pulveris smaller, never extending to the apex of basoflagellomere (Fig. 2A, C )); posterior surstyle lobe narrower and more elongated (Figs 8A :p, B:p, 10D) (in M. natans and M. pulveris posterior surstyle lobe broader (Fig. 10A-C )); ventral margin of posterior surstyle lobe with distinct triangular prominence (Figs 8A :v, B:v, 10D:v), which is not visible from lateral view in M. natans (Fig. 10A, B ); pile on terga longer, erect, predominantly pale, some black adpressed pile may present near posterior margin of terga 2-3 (in M. natans and M. pulveris pile on terga shorter, semi adpressed, black pile present in anterior and posterior part of terga 2-3). Females of M. makrisi sp. nov., M. natans and M. pulveris are very similar, but fossette position is near to the base of arista in M. makrisi sp. nov. (Fig. 2F, I :f) can differentiate this species from M. pulveris and M. natans (Fig. 2B, D, J :f, K:f) (in M. natans and M. pulveris fossette position medially between the base of the arista and apex of basofagellomere). Description. Male (Fig. 7A ). Head : Antenna : dark brown; basal part of arista light brown; in some specimens basoflagellomere ventrally light brown; basoflagellomere two times as long as wide with acute apex; large fossette extending from base of arista to apex of basoflagellomere (Fig. 2E ); lunule dark brown. Face : Black, white pollinose; covered with long white pile as long as pedicel, except for medial vitta extending from base of antenna to lower part of face without long white pile; ventral part of face and anteroventral part of gena black, shiny; frontal triangle black, white pollinose, covered with dense long white pile as long as pedicel; eyes holoptic, covered with dense white pile; in specimens from Israel and Somalia eyes covered with scarce, white pile as long as scapus; eye contiguity about 10 ommatidia long; vertical triangle isosceles black, shiny except for anterior part to anterior ocellus and posterior part to posterior ocelli white pollinose; vertical triangle covered with long white pile as long as pile on frontal triangle and intermixed black pile on ocellar triangle; ocellar triangle equilateral; occiput white pollinose covered with white pile as long as pile on vertical triangle. Thorax : Scutum black golden-bronze lustered with five white pollinose vittae; covered with yellow erect pile as long as pile on occiput and short black adpressed pile between transverse suture and scutellum; short black bristles present on area beyond transverse suture and above wing base; scutellum black, covered with long white pile as long as pile on scutum; pleura black, white pollinose; dorsal part of anterior anepisternum, posterior anepisternum, anterior anepimeron, dorsomedial part of anepimeron with long dense white pile as long as pile on scutum; long white pile on katepisternum broadly separated with bare area between; proepimeron and katatergum with some white pile. Legs : Femora black, yellow only at base and apex, covered with white pile; metafemur swollen with serrated triangular lamina (Fig. 4E ); tibiae of pro-, meso- and metaleg reddish-brown, medially darkened, covered with white pile; colour of basitarsomere, second and third tarsomeres of pro-, meso- and metaleg varies from yellow to dark brown; fourth and fifth tarsomeres always darkened - light to dark brown; tarsomeres mainly covered with white pile, intermixed with some black pile. Wing : Hyaline, covered with microtrichia except for some bare areas in first and second basal cells; stigma light yellow; wing veins dark brown, basally yellowish; halter and calypter yellow. Abdomen : Black with golden-bronze luster, slightly tapering; tergum 2 without antero-lateral reddish maculae; terga 2-4 with white pollinose fasciae; pollinose fascia on tergum 2 widely separated; lateral margin of abdomen blackish and white pollinose, posterior margin of terga 3-4 broadly yellowish; terga covered with long golden erect pile as long as pile on mesonotum and some short, black, adpressed pile may be present near posterior margin of terga 2-3; sternum 1 dark brown to blackish; colour of sterna 2-4 varies from dark brown to light brown; sterna covered with long, white, erect pile longer than pile on terga. Male genitalia (Figs 8 , 10D ): Epandrium (Fig. 8A :a, p, B:a, p): anterior surstyle lobe on inner side oval shaped, covered with dense, short, white pile (Fig. 8A :a, B:a); ventral margin of posterior surstyle lobe with distinct triangular prominence (Figs 8A :v, B:v, 10D:v); cercus square-like, in some specimens rounded (Fig. 8A :c). Hypandrium (Fig. 8C, E ): Broad medially (Fig. 8C ); aedeagus and associated structures as in Fig. 8D : ae, la, aa, ea, n; lateral sclerite of aedeagus elongate with small protuberance in posteroventral part, covered with short, white dense microtrichia (Fig. 8D : la). - Female (Fig. 7B ): Differs from male by normal sexual dimorphism and following characters: colours clearly different on mesoscutum and abdomen, golden luster lacks (Fig. 7A, B ); body pilosity distinctly shorter (Fig. 7A, B ); fossette smaller, never extending to apex of basoflagellomere (Fig. 2F ); frons black, white pollinose except shiny ocellar triangle and medial vitta extending from anterior part of anterior ocellus to lunule; pile on terga shorter and adpressed; terga 3-4 with short black adpressed pile on anterior and posterior parts. Variability. The population of Merodon makrisi sp. nov. in Cyprus is characterized by some stable morphological characters: eyes with dense white pile; tergum 2 without anterolateral reddish maculae; lateral margin of abdomen blackish; fasciae on terga smaller and narrower than in mainland specimens of M. makrisi sp. nov. and Autumn generation specimens of M. pulveris , while in specimens from mainland (Israel) these features are variable. The population of M. makrisi nov. from Cyprus shares very similar characteristics of its fasciae with the spring generation of M. pulveris : their smaller and narrower fasciae on terga, medially pointed at least on tergum 3, not reaching the lateral margin of terga are present also in the spring generation of M. pulveris . Etymology. The specific epithet is derived from the personal name Makris (a noun in genitive case). The species is dedicated to Christodoulos Makris, who collected the main part of the type series of the new species. He is an excellent observer, ecologist, photographer and author of Cyprus wildlife, insects and plants. Distribution. Merodon makrisi sp. nov. is found in Cyprus, Israel and Somalia (Fig. 6 ). Biology. In Cyprus, this species is commonly found on or near calcareous soils in rocky habitats with bulbs and orchids, around patches with open vegetation (Fig. 7C ). Besides Prospero autumnale , flowers and plants such as Narcissus sp., Cistus sp., Colchicum troodi Kotschy, Juniperus sp., Pistacia sp., Asphodelus ramosus L., Drimia aphylla (Forssk.) J.C. Manning & Goldblatt are found here, as well as Pinus brutia Ten. as the dominant tree. The fly is most abundant at lower altitudes up to about 200 m a.s.l., but also found higher up in the mountains. Flies found higher up (740 m a.s.l.), fly around swiftly, and often rest on low vegetation or on the ground. Flower visits have been observed on Prospero autumnale . Flight period is in Autumn (October/November). The puparium was found in Prospero autumnale and described here. On 7 Feb. 2016 bulbs of P. autumnale were collected (Fig. 7C ) and reared at ambient temperature. One adult male emerged on 21 Oct. 2016. The puparium remained in the bulb (Fig. 9A, B ). Figure 8. Merodon makrisi Vujic , Radenkovic & Tot sp. nov. male genitalia, Cyprus. A epandrium B left surstyle C , E hypandrium D aedeagus and associated structures; A , C , D lateral view B , E ventral view. Abbreviations: a - anterior surstyle lobe, aa - aedeagal apodeme, ae - aedeagus, c - cercus, ea - ejaculatory apodeme, la - lateral sclerite of aedeagus, n - notch on theca to which is attached the aedeagal apodeme, p - posterior surstyle lobe, v - triangular prominence on ventral margin of posterior surstyle lobe. Scale bar: 0.2 mm. Figure 9. Bulb of Prospero autumnale ( A , B ), from which one male specimen of Merodon makrisi Vujic , Radenkovic & Tot sp. nov. was reared. Photo: A. van Eck. Figure 10. Epandrium, lateral view. A , B Merodon natans C M. pulveris D M. makrisi Vujic , Radenkovic & Tot sp. nov. Abbreviation: v - triangular prominence on ventral margin of posterior surstyle lobe; A Greece (Achaia) B Montenegro C Greece (Rhodes) D Cyprus. Scale bar: 0.2 mm.