Research 75 (2017) 141e145

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Cretaceous Research

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Short communication First record of the subfamily Archinemestriinae in the family (Diptera: ) from Upper Cretaceous

* 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 Sciences and Technology of China, Hefei 230026, 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: Mesonemestrius caii gen. et sp. nov. is the first record of Archinemestriinae (Nemestrinidae) to be Received 13 October 2016 described from Upper Cretaceous Burmese amber. Mesonemestrius caii can be attributed to the subfamily Received in revised form Archinemestriinae by the following characters: M1 ending posterior to the wing tip and supernumerary 15 February 2017 crossveins absent. Mesonemestrius is characterized by M bifurcating after the conjunction of M þ þ and Accepted in revised form 3 March 2017 1 1 2 3 a long straight R . The discovery not only adds to the diversity of tangle-veined flies in Burmese amber, Available online 6 March 2017 1 but also represents the youngest record of the ancient subfamily Archinemestriinae. © 2017 Elsevier Ltd. All rights reserved. Keywords: Diptera Nemestrinidae Archinemestriinae Burmese amber Upper Cretaceous

1. Introduction 1960; Bernardi, 1973; Richter, 1997). Adult nemestrinids with highly developed flying abilities frequently visit flowers, often with The family Nemestrinidae, commonly called tangle-veined flies, an elongated proboscis as an adaptation to feeding on nectar is a quite small group of brachycerous Diptera with about 300 (Karolyi et al., 2013), but those with vestigial mouthparts do not extant and fossil in about 30 genera (Bernardi, 1973; feed (Teskey, 1981). Evenhuis, 1994; Mostovski and Martínez-Delclos, 2000; Brown Fossil nemestrinids are well recorded since the Mesozoic and et al., 2009). It is a cosmopolitan group, but uncommon in some probably originated in the Late /Early (Ansorge and regions (Gonzalez and Carvacho, 2016). The adult flies are medium- Mostovski, 2000). The most ancient and undoubted nemestrinid fly large, stout-body Diptera, often with a conspicuous dense pilosity; is found in the Lower Jurassic of Germany, and nemestrinids have the wings are usually longer than the body, venation complex with been abundant since the MiddleLate Jurassic with many species a compound diagonal vein, tibiae without apical spurs, and tarsi discovered in the of Kazakhstan, Zaza For- with an empodium (Yeates, 1994; Wedmann, 2007). mation of Russia, of China and La Pedrera outcrop The biology of Nemestrinidae is moderately well known: the of Spain and three species (two new) described from Upper larvae of all known species are internal , they parasitize Cretaceous Burmese amber (Rohdendorf, 1968; Mostovski, 1998; the egg clusters and adult bodies of other , such as scar- Ren, 1998; Mostovski and Martínez-Delclos, 2000; Grimaldi, abaeid beetles and acridid Ortheoptera (Greathead, 1958; Prescott, 2016). The subfamily Archinemestriinae Rohdendorf, 1968 is an extinct subfamily that is only found in the Upper Jurassic of Kazakhstan: here we report a new species and in Archine- * Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, mestriinae from Upper Cretaceous Burmese amber, that is also the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 youngest record of this subfamily. East Beijing Rd., Nanjing 210008, China. E-mail address: [email protected] (B. Wang). http://dx.doi.org/10.1016/j.cretres.2017.03.005 0195-6671/© 2017 Elsevier Ltd. All rights reserved. 142 Q. Zhang et al. / Cretaceous Research 75 (2017) 141e145

2. Material and methods Mesonemestrius caii sp. nov. (urn:lsid:zoobank.org:act:AFB364B2-C531-400D-931B- The specimen described herein originated from the Hukawng 1D93F140ABFF) Valley of Kachin Province, Myanmar (locality in Kania et al., 2015: Figs. 1e5 fig 1). The record of Cretaceous insects has increased dramatically Derivation of name. The specific name caii is in honor of Mr. Cai during the past twenty years due to discoveries of many new fossils Yong, for his donation of this specimen. including compressions and amber (Wang and Szwedo, 2015). Description. Based on female. A small-sized fly with body length Burmese amber, which is considered as earliest in age, 4.02 mm; head length 0.51 mm, width 1.29 mm; thorax length has a very high diversity of inclusions: about 36 orders and over 1.27 mm, width 1.10 mm; wing length 4.42 mm, width 1.37 mm; 250 families of are known; as well as invertebrates, halter length 0.66 mm; abdomen length 1.96 mm (excluding cerci), protists, plants, fungi and vertebrate inclusions are known (Ross widest segment of abdomen 1.11 mm across; cerci 0.28 mm long. et al., 2010; Shi et al., 2012). The amber piece has been polished with sand paper with Head hemispherical, occiput flat; eyes large, separated by a different grain sizes and with polishing power, the crack in amber narrow frons; antenna with scape cylindrical, pedicel subspherical has been bonded with UV glue in full and then irradiated by ul- and shorter than scape, flagellum with four slender flagellmores, traviolet lamp. The amber is housed in the Nanjing Institute of about 4 times longer than scape and pedicel combined; flag- Geology and Palaeontology (NIGP), Chinese Academy of Sciences. ellomere 1 is thickest and longest, flagellomere 2 and flagellomere Observations were made using an Olympus SZX7 Microscope and 3 slightly decending, flagellomere 4 tapered in width apically; photographs were taken using a Zeiss Stereo Discovery V16 mi- ocellar tubercles subtrianglular, ocelli not recognizable; mouth- croscope system and Confocal Laser Scanning Microscope (CLSM), parts with only labellum visible, labella flabelliform (Fig. 2). the photomicrographs were taken using a CLSM Zeiss LSM710 Thorax narrower than head, scutellum triangular; wings narrow with 10 objectives and a laser at 488 nm. In most instances, and long, with alula and anal lobe vestigial; costal vein thick, incident and transmitted light were used simultaneously. In the becoming thinner beyond wing tip; costal margin of subcostal cell case of some reflected light caused by convex surfaces, we dropped distinctly longer than that of first radial cell, about 2 times longer; some glycerol on a cover glass, and then deposited the glass on the R1 about nine-tenths of wing length; R ending just behind wing amber. All images are digitally stacked photomicrographic com- 5 tip; dR þ forked distinctly basad of R apex; R þ and M þ fused posites of approximately 20 individual focal planes obtained using 4 5 1 4 5 1 2 for some distance, R þ þM þ as long as bR þ and dR þ ; vein r-m the free software Combine ZP for a better illustration of the 3D 4 5 1 2 4 5 4 5 missing; diagonal vein long and straight, composed of Rs, bR þ , structures. The figures were prepared with CorelDraw X7 and 4 5 R þ þM þ ,dMþ ,Mþ þ ,Mþ ,dM and dM þ ;Mþ stem Adobe Photoshop CS6. Vein abbreviations follow Zhang (2010). All 4 5 1 2 1 2 1 2 3 2 3 3 3 4 2 3 directly continuous with dM ; crossvein m-cu connecting CuA to taxonomic acts established in the present work have been regis- 3 M ; CuA and CuP subparallel; cell cu (traditionally anal cell) widely tered in ZooBank (see below), together with the electronic publi- 4 open, anula degraded, anal lobe narrow; supernumerary crossveins cation LSIC: urn:lsid:zoobank.org:pub:BDACBFE6-7DA5-4411- absent (Fig. 3); halter relatively long, with a dark club and pale 902B-68FEFC8237F8. stalk, club obtuse; legs simple and slender, foreleg and midleg relatively short, about 20 mm long; hindleg relatively long, about 2 times longer than foreleg and midleg; tibial spurs absent; pulvilli- 3. Systematic paleontology form empodium well developed; first tarsomere of foreleg 0.58 mm long, slightly longer than other four combined; ratio of protarso- Order Diptera Linnaeus, 1758 meres 3.6: 1.0: 0.5: 0.4: 1.4; first tarsomere of midleg 0.42 mm long, Suborder Brachycera Zetterstedt, 1842 relatively longer than other four combined, ratio of meso- Family Nemestrinidae Macquart, 1834 tarsomeres 7.0: 1.0: 1.0: 0.66: 1.66; first tarsomere of hindleg Subfamily Archinemestriinae Rohdendorf, 1968 0.52 mm long, relatively longer than other four combined; ratio of Genus Mesonemestrius gen. nov. metatarsomeres 4.0: 1.0: 0.7: 0.5: 0.9 (Fig. 4). (urn:lsid:zoobank.org:act:EB12123A-9B70-403D-8ABA- Abdomen with ten tergites; first tergite quite short, third tergite 9D882728997F) longest, fourth tergite widest; width of third to eighth tergites decreasing, and ninth tergite thin and long, cylindrical, tenth tergite Derivation of name. The generic epithet is derived from Mesozoic quite short, ninth and tenth tergites as wide as the base of cerci; and known genera Archinemestrius and Protonemestrius in the length of first to ninth tergites: 0.09 mm, 0.26 mm, 0.45 mm, subfamily Archinemestriinae. 0.34 mm, 0.26 mm, 0.12 mm, 0.10 mm, 0.08 mm, 0.13 mm and Type and only species. Mesonemestrius caii sp. nov. 0.05 mm, respectively; spermathecae invisible; cerci length Material. Holotype specimen number NIGP165265 (female); stored 0.28 mm, saber-shaped, 3 times as long as wide at base; cerci at the Nanjing Institute of Geology and Palaeontology (NIGP), Chi- distinctly backward curved except base; cerci seem slightly scler- nese Academy of Sciences, Nanjing, China. otized and setulous; terminal sternite notched with fine setae, Diagnosis. Small-sized flies, head hemispherical, a little broader setae around notch slightly longer (Fig. 5). than thorax; costal vein becoming thinner before wing tip; costal margin of subcostal cell distinctly longer than that of first radial cell; R1 very long and straight; R5 ending just behind wing tip; 4. Discussion R4þ5 and M1þ2 fused for some distance, crossvein r-m missing; diagonal vein long and straight, composed of Rs, bR4þ5, Rohdendorf (1968) erected the subfamily Archinemestriinae R4þ5þM1þ2,dM1þ2,M1þ2þ3,M2þ3,dM3 and dM3þ4; anal cell for two genera from the Jurassic of Karatau (Kazakhstan), based widely open; anula and anal lobe vestigial; supernumerary especially on the phragma and M1 ending posterior to the wing crossveins absent; antenna with 4 flagellomeres, tapered apically; tip. Mostovski (1998) revised the diagnosis of the subfamily first tarsomere of midleg longer than other four combined; cerci Archinemestriinae and described four new species in two genera, relatively thin and long, terminal sternite notched; mouthparts Archinemestrius and Protonemestrius, from the same locality. The with labellum flabelliform. new genus can be placed in the subfamily Archinemestriinae Q. Zhang et al. / Cretaceous Research 75 (2017) 141e145 143

Fig. 3. Mesonemestrius caii gen. et sp. nov., holotype NIGP165265. A, Microphotograph of wing; B, Line drawings of wing. (Scale bars represent 0.5 mm).

based on the following characters: M1 ending behind the wing tip and supernumerary crossveins absent. Mesonemestrius differs from Archinemestrius in the costal margin of the subcostal cell being distinctly longer than that of the first radical cell; from Protonemestrius in mouthparts being vestigial, R4þ5 and M1þ2 fused for a rather long distance, and cerci thin and long. Apart from the diagnosis of the subfamily Archinemestriinae, one feature occurs in all known species of Archinemestriinae: R1 is relatively short and curved upwards terminally, but Meso- nemestrius has R1 very straight and long, sharing a similar appearance with extant nemestrinids flies. Mesonemestrius is also characterized by M1 bifurcating after the conjunction of M1þ2þ3. Mesonemestrius shares more apomorphic and plesiomorphic characters with Archinemestrius (R5 ending behind the wing tip, first tarsomere of the midleg being longer than the other four combined, cerci relatively long and narrow) and has more advanced characteristics than Archinemestrius (R1 long and Fig. 1. Mesonemestrius caii gen. et sp. nov., holotype NIGP165265. A, Microphotograph straight, M1þ2 fused with M3 for some distance, cerci backward of dorsal features; B, Microphotograph of ventral view. (Scale bars represent 1.0 mm). curved and slightly sclerotized). Grimaldi (2016) described three species (two new) in the extant genus Hirmoneura from the same locality as Mesonemestrius caii. The wing vernation and cerci of M. caii are quite similar to H. zigrasi, but M. caii differs from H. zigrasi in having a much smaller size, the length of whole body only 4.02 mm, (H. zigrasi with thorax þ abdomen has a length of 8.5 mm); the cerci are also much thin and long, 3 times as long as wide (H. zigrasi only 2.2 times); alula and anal lobe vestigial (H. zigrasi has anal lobe absent but alula present); R5 and M1 terminating behind wing tip (H. zigrasi with R5 just before wing tip); M1 and M2 parallel to R5 (H. zigrasi with M1 and M2 upcurved, a little convergent to wing tip). Based on M1 ending behind wing tip and comparison with M. caii, H. zigrasi might belong to the extinct subfamily Archinemestriinae. There are many Mesozoic records of nemestrinid flies, but nemestrinids being reported in amber is generally quite unusual; till now, only three species have been described from amber (Grimaldi, 2016). This may be because they are often large and fast fliers, and hardly ever get caught in resin and if so, can easily break out; another reason may be that they prefer open habitats rather than forested ones where resin is found (Kosmann, 2015). The Fig. 2. Mesonemestrius caii gen. et sp. nov., holotype NIGP165265. A, Microphotograph Archinemestriinae is a primitive group in the family Nemestrinidae, of head in dorsal view; B, Microphotograph of head in ventral view; C, Microphoto- graph of antenna; D, Line drawings of antenna. (Scale bars represent 0.2 mm). the earliest and only previously known members of this extinct 144 Q. Zhang et al. / Cretaceous Research 75 (2017) 141e145

also the first three-dimensionally preserved archinemestriin specimen and the fourth record of the family Nemestrinidae in amber. Bernardi (1973) completed the subfamily classification of Nemestrinidae: Bequaert (1930, 1932) recognized three subfamilies whereas Sack (1933) only two, Greathead (1967) three, accepting the system proposed by Bequaert, but with special attention to the male genitalia; Rohdendorf (1968) erected Archinemestriinae (above) and Bernardi (1973) proposed a classification with 7 different subfamilies, with two new subfamilies and a genus incertae sedis, but this was not generally accepted. The classification of the subfamilies in Nemestrinidae is therefore quite uncertain, but the subfamily Archinemestriinae Rohdendorf (1968) is un- doubtedly an extinct subfamily of Nemestrinidae. Although the general subfamily classification is uncertain, a definite synapomorphic character of all known genera of Nem- estrinidae is that the diagonal vein starts from cell br and the 4th posterior cell is closed well before the wing margin (Nagatomi and Yang, 1998). Sinonemestrius tuanwangennsis Hong & Wang, 1990 was described from the Lower Cretaceous Laiyang Forma- tion in Shandong Province, China, and was originally placed in Nemestrinidae, but Mostovski and Martínez-Delclos, 2000 and Jarzembowski and Mostovski (2000) thought that the characters including the short vein R1 andarchedR2þ3 terminating at the tip of R1 and absence of diagonal vein show that this genus does not belong in Nemestrinidae. Zhang et al. (2008) described Ahirmoneura neimengguensis fromtheMiddleJurassicofInner Mongolia, China. Ahirmoneura has some similarities with Sino- nemestrius Hong & Wang, 1990, and its wing venation lacking diagonal veins, crossvein r-m located at the extreme base of cell d, cell m3 ending before cell d shows that it cannot be attributed to Nemestrinidae.

5. Conclusion

This paper describes a new tangle-veined fly Mesonemestrius caii gen. et sp. nov. from Upper Cretaceous Burmese amber. It represents the first record of the subfamily Archinemestriinae in the family Nemestrinidae in Burmese amber and is the latest occurrence of the subfamily Archinemestriinae. Our find increases the diversity of brachycerous flies and Nemestrinidae during the Late Cretaceous, and is the first paper with detailed characters of Fig. 4. Mesonemestrius caii gen. et sp. nov., holotype NIGP165265. A, Microphotograph the fossil antenna and cerci in the subfamily Archinemestriinae. of foreleg; B, Line drawings of protarsomeres; C, Microphotograph of midleg; D, Line drawings of mesotarsomeres; E, Microphotograph of hindleg; F, Line drawings of metatarsomeres. (Scale bars represent 0.2 mm). Acknowledgements

We are grateful to Neal Evenhuis and one anonymous reviewer for their useful comments on the first version of the paper. We also thank E.A. Jarzembowski for improving the English language of the manuscript. This research was supported by the National Natural Science Foundation of China (41572010, 41622201, 41688103), and Youth Innovation Promotion Association of CAS (No. 2011224).

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