Cretaceous Research 67 (2016) 1e7

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Cretaceous Research

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A remarkable brachyceran fly (Diptera: Tabanomorpha) from Late Cretaceous Burmese amber

* Qingqing Zhang a, b, Junfeng Zhang a, c, Bo Wang a, d, a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Rd., Nanjing 210008, China b University of the Chinese Academy of Sciences, Beijing 100049, China c College of Palaeontology, Shenyang Normal University, Shenyang 110034, China d Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Science, Beijing 100101, China article info abstract

Article history: A new brachyceran fly (Diptera: Tabanomorpha), Pseudorhagio zhangi gen. et sp. nov., is described from Received 6 April 2016 Late Cretaceous Burmese amber. It is tentatively placed as Family incertae sedis in Tabanomorpha and Received in revised form distinguished from other Tabanomorpha by the following unusual combination of characters: head wider 22 June 2016 than thorax; body densely covered with fine and short setae, devoid of macrosetae; scutum strongly Accepted in revised form 26 June 2016 convex, nearly spherical; scutellum rather small, convex; antennal flagellum elongated, tapering, un- Available online 27 June 2016 segmented; vein R4 perpendicular to R5, and strongly curved; crossvein m-m very long, strongly sinu- ated; tibial spur formula 0, 2, 0. This discovery further confirms the high diversity of Tabanomorpha in Keywords: Insecta Late Cretaceous Burmese amber. An updated list of brachyceran species in Burmese amber is given. © Diptera 2016 Elsevier Ltd. All rights reserved. Tabanomorpha Burmese amber Cretaceous

1. Introduction following apomorphic characters: adult with convex and bulbous clypeus; expanded first article of the female cercus; a brush on the The Diptera represent an order that is one of the most larval mandible and larval head retractile (Yeates and Wiegmann, species-abundant, ecologically diverse and evolutionarily signifi- 1999; Yeates, 2002; Santos, 2008; Kerr, 2010). Until now, a large cant groups. The order Diptera is composed of two suborders: number of Mesozoic Tabanomorpha have been described world- and Brachycera. Brachycera are composed of four wide, including more than 80 rhagionid species in more than 40 infraorders, the lower Brachycera: Xylophagomorpha Fallen, 1815, genera (Zhang, 2010, 2011, 2013; Nel et al., 2014; Angelini et al., Stratiomyomorpha Hennig, 1973, Tabanomorpha Hennig, 1948 2016), seven tabanid species in five genera (Zhang, 2012), and 12 and the higher Brachycera: Sharp, 1894 (Yeates athericid species in seven genera (Zhang, 2012; Oberprieler and and Wiegmann, 1999; Wiegmann et al., 2000; Yeates, 2002). Yeates, 2014), and to date, no record of vermileomid and peleco- Krzeminski and Krzeminska (2003) added rhynchid flies have been found as Mesozoic fossils. Rohdendorf, 1961 in lower Brachycera. Tabanidae Latreille, 1802, Among the insect taxa preserved in amber, dipterans are the Enderlein, 1922, Latreille, 1802, most common and diverse organismal inclusions (Grimaldi and Stuckenberg, 1973, and Nagatomi, 1977 Cumming, 1999; Grimaldi et al., 2002; Dikow and Grimaldi, have been commonly assigned to Tabanomorpha based on the 2014). Many brachyceran flies as amber inclusions have been studied in Lebanese, Canadian, Siberian, French, New Jersey, Spanish and Burmese ambers (Grimaldi and Cumming, 1999; Perrichot, 2004; Grimaldi and Arillo, 2008; Grimaldi et al., 2009, 2011; Dikow and Grimaldi, 2014; Arillo et al., 2015; Angelini * Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, et al., 2016; Zhang et al., 2016). Several brachyceran flies have Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 been described from Burmese amber (see Table 1), and here we East Beijing Rd., Nanjing 210008, China. E-mail address: [email protected] (B. Wang). describe a new genus and species within lower Brachycera in http://dx.doi.org/10.1016/j.cretres.2016.06.012 0195-6671/© 2016 Elsevier Ltd. All rights reserved. 2 Q. Zhang et al. / Cretaceous Research 67 (2016) 1e7

Table 1 List of brachyceran species in Burmese amber, modified and updated from Grimaldi et al., 2009, 2011; Arillo et al., 2015; Dikow and Grimaldi 2014.

Infraorder Family Genus Species

Stratiomyomorpha Zhangsolvidae Nagatomi and Yang, 1998 Linguatormyia Grimaldi, 2015 Linguatormyia teletacta Grimaldi, 2015 Eremochaetidae Ussatchev, 1968 Zhenia Zhang et al., 2016 Zhenia xiai Zhang et al., 2016

Muscomorpha Chimeromyiidae Grimaldi and Cumming, 2009 Chimeromyia Grimaldi and Cumming, 1999 Chimeromyia burmitica Grimaldi & Cumming, 2009 Leach, 1815 Schlingeromyia Grimaldi & Hauser, 2011 Schlingeromyia minuta Grimaldi & Hauser, 2011 Burmacyrtus Grimaldi & Hauser, 2011 Burmacyrtus rusmithi Grimaldi & Hauser, 2011 Zaitzev, 1991 Microburmyia Grimaldi & Cumming, 2011 Microburmyia analvena Grimaldi & Cumming, 2011 Microburmyia veanalvena Grimaldi & Cumming, 2011 Nagatomi, 1991 Kumaromyia Grimaldi & Hauser, 2011 Kumaromyia burmitica Grimaldi & Hauser, 2011 Apystomyiidae Nagatomi and Liu, 1994 Hilarimorphites Grimaldi & Cumming, 1999 Hilarimorphites burmanica Grimaldi & Cumming, 2011 Tethepomyiidae Grimaldi and Arillo, 2008 Tethepomyia Grimaldi & Cumming 1999 Tethepomyia zigrasi Grimaldi & Arillo, 2011 Unplaced Myanmyia Grimaldi, 2011 Myanmyia asteiformia Grimaldi, 2011 Latreille, 1802 Burmapogon Dikow & Grimaldi, 2014 Burmapogon bruckschi Dikow & Grimaldi, 2014

Tabanomorpha from Burmese amber based on two relatively well dated at 98.79 ± 0.62 Ma based on UePb zircon dating of the vol- preserved specimens. canoclastic matrix (Shi et al., 2012). The specimens are preserved in two pieces of yellow amber with some impurities. The amber pieces 2. Material and methods containing the inclusions were cut, trimmed and polished. The specimens are housed in the Nanjing Institute of Geology and The specimens described herein were collected from the Palaeontology (NIGP), Chinese Academy of Sciences. Photographs Hukawng Valley of Kachin Province, Myanmar (for locality see Kania were taken using a Zeiss Stereo Discovery V16 microscope system et al., 2015: fig. 1). The age of Burmese amber is radiometrically and Zen software. In most instances, incident and transmitted light

Fig. 1. Pseudorhagio zhangi gen. et sp. nov. A, Photograph of holotype, NIGP164490, in dorsal view; B, Photograph of paratype, NIGP164491, in lateral view; C, Photograph of paratype, NIGP164491, in lateral view; D, Drawing of paratype, NIGP164491. Q. Zhang et al. / Cretaceous Research 67 (2016) 1e7 3

Fig. 2. Pseudorhagio zhangi gen. et sp. nov. A, Photograph of holotype, NIGP164490, head; B, Photograph of paratype, NIGP164491, antenna; C, Drawing of holotype, NIGP164490, antenna; D, Drawing of paratype, NIGP164491, antenna. were used simultaneously. All images are digitally stacked photo- Type species: Pseudorhagio zhangi sp. nov. micrographic composites of approximately 20 individual focal Diagnosis. Head semispherical, approximately the same width as planes obtained using the free software Combine ZP for a better thorax; antennal flagellum elongate-subuliform, unsegmented; illustration of the 3D structures. The figures were prepared with scutum strongly convex, nearly spherical; scutellum very small, CorelDraw X7 and Adobe Photoshop CS3. All taxonomic acts concave; costal vein circumambient, weakened beyond vein R4; established in the present work have been registered in ZooBank R4þ5 fork shallow but wide, distinctly distal to end of cell d; M (see below), together with the electronic publication LSID: urn:l- with three branches; crossvein m-m elongate, strongly sinuated; sid:zoobank.org:pub:5EF0E953-92E8-461E-8B0E-66482FFC8FC5. discal cell narrow and long, drift apically; cell cu narrowly open; anal lobe distinct; tibial spur formula 0, 2, 0. 3. Systematic palaeontology Pseudorhagio zhangi sp. nov. (urn:lsid:zoobank.org:act:87489E62-F82F-45FA-9CD0- Order Diptera Linnaeus, 1758 90E1BE52DFB6) Suborder Brachycera Zetterstedt, 1842 Figs. 1e4 Infraorder Tabanomorpha Hennig, 1948 Family Unknown Derivation of name. The specific name is in honour of Prof. Haichun Zhang, palaeoentomologist. Genus Pseudorhagio gen. nov. Material. Holotype NIGP164490, Paratype. NIGP164491. (urn:lsid:zoobank.org:act:4FBA9976-68DB-4735-91F9- Diagnosis. R4 strongly curved, perpendicular to R5 at base and AE558D6FB5CA) terminating just before wing apex; m-m strongly sinuated, Derivation of name. The generic name is derived from the Greek perpendicular to M4. ‘Pseudo’ and the genus ‘Rhagio’ (rhagionid fly). Description. Based on two adult flies. 4 Q. Zhang et al. / Cretaceous Research 67 (2016) 1e7

Fig. 3. Pseudorhagio zhangi gen. et sp. nov. A, Photograph of holotype, NIGP164490, left wing; B, Drawing of holotype, NIGP164490, left wing; C, Photograph of paratype, NIGP164491, left wing. Q. Zhang et al. / Cretaceous Research 67 (2016) 1e7 5

Fig. 4. Pseudorhagio zhangi gen. et sp. nov., holotype NIGP164490. A, Photograph of fore tarsus; B, Photograph of mid tarsus; C, Photograph of hind tarsus.

Body length 3.8e4.6 mm; wing length 3.3e3.6 mm, width crossvein m-cu as long as crossvein r-m; cell br slightly longer and 1.3e1.5 mm; length of tibia: fore 0.9 mm, mid 1.1 mm, hind 1.0 mm wider than cell bm; apical part of CuA weakly arched; CuP almost in holotype. straight; tips of CuA, CuP ending at hind margin, cell cu narrowly Head very large, 0.7 mm long, 1.1 mm wide, slightly wider than open (Fig. 3A, B); lower calypter reduced, indistinct in paratype thorax (1.0 mm wide); eyes large, dichoptic in paratype; antenna (Fig. 3C); halter short, stout, haltere long in paratype. with scape cylindrical, pedicel subspherical and distinctly shorter Legs slender, long, covered with dense macrosetae; tibial spurs than scape, flagellum slender and long, tapering at apex, nearly 2 with following formula 0, 2, 0; empodium pulvilliform; pulvilli times longer than scape and pedicel combined (Fig. 2); mouthparts shorter than empodium; fore femur 0.87 mm long, tibia 0.92 mm with only labellum visible, labellum laminar, fringed with long long, tarsus 0.98 mm long; mid femur 1.06 mm long, tibia 1.14 mm setae in paratype. long, tarsus 1.17 mm long; hind femur 1.45 mm long, tibia 1.02 mm Thorax 1.1 mm long, 1.0 mm wide, about 1.5 mm deep, covered long, tarsus 1.33 mm long; ratio of fore tarsomeres 2.4: 1: 0.6: 0.4: densely with short setae; scutum very large, spherical, exceedingly 0.7, ratio of mid tarsomeres 2.6: 1: 0.7: 0.5: 0.4, ratio of hind tar- convex in paratype; scutellum rather small, triangular in paratype; someres 2.5: 1: 0.7: 0.3: 0.5 (Fig. 4). wing relatively long and wide, with length 3.3 mm, width 1.4 mm, Abdomen 2e2.2 mm long, 0.8 mm wide, subcylindrical but apex rather obvious, extending well beyond apex of abdomen; slightly tapered, some 3.5 times as long as head (excluding an- costal vein thick, covered with short setae, thinner beyond R4; tenna) and about 2 times longer than thorax; eight abdominal length of costal section R1eR2þ3 about two times of section Sc-R1, tergites visible, first tergite slightly shorter than remainder, length and distinctly shorter than section R2þ3eR4þ5; stem R4þ5 (i.e., of second to sixth tergites decreasing, seventh tergite slightly bR4þ5þ dR4þ5) about 3 times longer than stem Rs; R4þ5 1.6 times shorter than second tergite, eighth tergite distinctly narrower than longer than R5; distal cell narrow and long, 3.8 times as long as other tergites; all tergites densely setose, genitalia structures been wide; crossvein r-m long, about one half of length of bR4þ5 and sheltered by hairs on eighth tergite. dividing anterior margin of discal cell 1:2; M1 faintly arched anteriorly and sharply down curved posteriorly; bM2 and dM2 4. Discussion straight forming a right angle; dM3þ4 straight, subparallel to dM2; costal section M1eM2 as long as costal section M2eM3þ4, distal of Pseudorhagio zhangi can be placed in Tabanomorpha based on m-m strongly curved, formed a right angle with base of M3þ4; the following characters: stout body, bulbous clypeus, vein R1 and 6 Q. Zhang et al. / Cretaceous Research 67 (2016) 1e7

R2þ3 relatively long, cell d relatively large, and all posterior cell continued discovery and documentation of Cretaceous Tabano- open. According to the key to tabanomorphan families provided by morpha can help build robust analyses for the phylogeny of Taba- Yeates (2002), it differs from Athericidae in having the antennal nomorpha and even Diptera. flagellum tapering at the apex; additionally, it has different wing venation according to Mostovski et al. (2003) and Zhang (2012): Acknowledgements cell r1 not closed and R4 and R5 diverging at right angles. P. zhangi is probably related to Tabanidae in R4 and R5 being divergent, cell This research was supported by the National Basic Research r4 relatively short and broad, CuP almost straight and nearly fused Program of China (2012CB821900), National Natural Science distally with CuA (Colless and McAlpine, 1991), but differs from this Foundation of China (41572010), and Youth Innovation Promotion family in having the lower calypter reduced and R5 not so far Association of the Chinese Academy of Sciences (No. 2011224). We behind the wing apex (Yeates, 2002; Kerr, 2010). On the other hand, are grateful to two anonymous reviewers for a critical review and P. zhangi differs from Vermileonidae in having the wings broadened useful suggestions. at the base and without apical spurs on the front tibiae (Borror et al., 1989); from Pelecorhynchidae in having vein CuP straight, References convergent with CuA at the apex of the wing (Colless and McAlpine, 1991); from Rhagionidae in having the base of fork R4 and R5 distal Angelini, P., Azar, D., Nel, A., 2016. A new genus and species of snipe fly (Diptera: to d according to Grimaldi and Cumming (1999). Rhagionidae) in Lebanese Cretaceous amber. Cretaceous Research 58, 10e16. ~ P. zhangi probably shares some features with Rhagionidae, Arillo, A., Penalver, E., Perez De La Fuente, R., Delclos, X., Criscione, J., Barden, P.M., Riccio, M.L., Grimaldi, D.A., 2015. Long-proboscid brachyceran flies in Creta- because both families have some shared features in the wing ceous amber (Diptera: Stratiomyomorpha: Zhangsolvidae). Systematic Ento- venation and tibial spur formula. Grimaldi and Cumming (1999) mology 40, 242e267. noted a potential synapomorphy of most Rhagionidae as “a base Borror, D.J., Triplehorn, C.A., Johnson, N.F., 1989. Chapter 32, Order Diptera. In: An e Introduction to the Study of , sixth ed. Saunders College Publishing, of fork R4 R5 that is at the same level as the distal end of cell d, and Pennsylvania, pp. 499e575. not distal to it; vein R5 is almost always straight and R4 arises from Colless, D.H., McAlpine, D.K., 1991. Diptera. In: CSIRO (Ed.), The Insects of Australia, it with a sharp bend at its base, often at 90”. P. zhangi has the second ed., vol. II. Melbourne University Press, Melbourne, pp. 717e786. fl synapomorphy of vein R5 and R4 formed at right angles, but the Dikow, T., Grimaldi, A., 2014. 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