生命科学学院-王永杰.pdf
生命科学学院-王永杰.pdf
Zootaxa 4691 (2): 153–160 https://www.mapress.com/j/zt/ Copyright © 2019 Magnolia Press Article ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4691.2.4 http://zoobank.org/urn:lsid:zoobank.org:pub:84158919-975D-4118-B0FD-45449337902F A new fossil snipe fly with long proboscis from the Middle Jurassic of Inner Mongolia, China (Diptera: Rhagionidae) YE HAN, YAJING CAI, DONG REN & YONGJIE WANG* Key Lab of Insect Evolution and Environmental Changes, Capital Normal University, Beijing 100048, China *Corresponding author. E-mail: wangyjosmy@gmail.com Abstract A new genus and species of rhagionids with a long proboscis, Elliprhagio macrosiphonius gen. et sp. nov., is described from the Middle Jurassic Jiulongshan Formation in Daohugou, Inner Mongolia, China, which is considered to be the earliest hematophagous rhagionid described hitherto according to the typically piercing and sucking mouthparts. All previously documented rhagionids from northeastern China are reviewed a key to genera of Rhagionidae from Daohugou is provided for the first time. The genus Daohugorhagio Zhang, 2013 is considered as a new synonym of Trichorhagio Zhang, 2013. Key words: fossil, new species, new genus, Daohugou, hematophagous feeding Introduction Rhagionidae, as a relic family of lower Brachycera, have been in existence for more than 240 million years. The earliest rhagionid fossil hitherto, Gallia alsatica Krzemiński & Krzemińska, 2003, was described from the Arzviller, France (Lower/Middle Triassic) (Krzeminski & Krzeminska 2003). The rhagionids, commonly known as snipe flies, began a rapid radiation since the Triassic. Five species belonged to four genera were described from the Early Jurassic (Ansorge 1996; Mostovski & Jarzembowski; 2000; Krzemiński & Ansorge 2005). In the Middle Jurassic, Rhagionidae started to show species richness and morphological diversity (Kovalev 1981; Evenhuis 1994) with 21 species in 14 genera described so far. They comprise nine species within five genera from Transbaikalia, and 12 species within nine genera from Daohugou (Kovalev 1981, 1982; Kalugina & Kovalev 1985; Zhang K. et al. 2006, 2008; Zhang J. 2010, 2011, 2013; Zhang J. et al. 2012). In the Late Jurassic, the generic diversity of Rhagionidae decreased gradually. Up to date, 13 species in five genera have been reported, most of which are from the Karabastau Formation, except for Palaeoarthroteles Kovalev & Mostovski, 1997 from the Glushkovo and Godymboy Formations (Rohdendorf 1938, 1964; Kovalev 1982; Kovalev & Mostovski 1997; Mostovski 2000, 2008). Comparing to the relatively restricted localities in the Jurassic, the Cretaceous rhagionids had much broader distribution with 20 species referred to 14 genera described from Asia, Europe, North America and Australia (Handlirsch 1906; Jell & Duncan 1986; Kovalev 1986; Zaitzev 1986; Hong et al. 1992; Zhang J. et al. 1993; Ren 1998; Grimaldi & Cumming 1999; Mostovski et al. 2000; Kraemer & Nel 2009; Angelini et al. 2016). In the Cenozoic, although exhibiting a low generic diversity with only three described genera so far, rhagionids had a noticeable diversification across species with 22 species found from Baltic, America, Germany and France (Meunier 1899, 1902, 1910, 1916; Cockerell 1908, 1911, 1921; Théobald 1937; Statz 1940; Melander 1949; Evenhuis 1994; Kerr 2010; Nel et al. 2016). Up to date, 35 fossil genera including 80 species have been attributed to Rhagionidae. Although significant progress on the taxonomy of fossil rhagionids have been made, a comprehensive review of the fossil rhagionids is still essential and crucial. The Middle Jurassic Daohugou locality is well-known for its diversity and quantity of insect fossils (Gao et al. 2012; Gu et al. 2012; Wang et al. 2012; Yao et al. 2014), and so far, 12 species in nine genera of Rhagionidae have been described from this locality. Many rhagionids from this locality, having well-preserved complete body and other appendages, provide detailed morphological features for us. In this paper, we describe a new genus and species Accepted by V. Blagoderov: 2 Oct. 2019; published: 30 Oct. 2019 153 with a long proboscis, Elliprhagio macrosiphonius gen. et sp. nov. The well-preserved mouthpart structure infers that the new species was adapted to piercing and feeding on blood, which is proposed to be the earliest hematophagous rhagionid hitherto. In addition, a key to genera of fossil rhagionids from Daohugou is given. Material and methods The specimens in this study were examined dry or under alcohol using a Leica M165C dissecting microscope with an attached digital camera system and illustrated with the aid of a drawing tube attachment. Line drawings are made in Adobe Photoshop CC 2014 (64 bit). The type specimens are deposited in the fossil insect collection of the Key Lab of Insect Evolution & Environmental Changes, Capital Normal University, Beijing, China. Vein nomenclature used here follows Wootton & Ennos (1989). Systematic Paleontology Key to genera of rhagionids from the Daohugou locality (some questionable genera are omitted) 1 2 3 4 5 6 7 8 9 - R4 diverging from R5, forming a short and wide cell r4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 R4 nearly parallel to R5, forming a long and narrow cell r4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 M1 and M2 converging to a point. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archrysopilus Zhang, Yang & Ren, 2008 M1 and M2 nearly parallel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ussatchovia Rohdendorf, 1964 R2+3 curved at the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 R2+3 nearly straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 M3 absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palaeobolbomyia Kovalev, 1982 M3 present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 CuP and A1 converging to a point and with a short petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elliprhagio gen. nov. CuP and A1 converging. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protorhagio Rohdendorf, 1938 M3 parallel to M4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 M3 slightly diverging from M4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 d cell rather narrow and long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinorhagio Zhang, Yang & Ren, 2006 d cell wide and long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 bM3 much shorter than dM3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parachrysopilus Zhang, 2013 bM3 nearly as long as dM3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palaeoarthroteles Kovalev & Mostovski, 1997 M3 and M4 converging to a point. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lithorhagio Zhang, 2012 M3 and M4 converging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichorhagio Zhang, 2013 Order Diptera Linneaus, 1758 Suborder Brachycera Zetterstedt, 1842 Family Rhagionidae Latreille, 1802 Genus Elliprhagio gen. nov. urn:lsid:zoobank.org:act:18DC6062-EF6F-40E8-901B-733F227C0CEE Type species. Elliprhagio macrosiphonius sp. nov. Diagnosis. Flagellum with 10 flagellemeres; proboscis long, labium fleshy, labella small. Wings elliptic and wide; R2+3 sinuate at the middle, and sharply up-curved distally; crossvein r-m intersecting the upper margin of d cell at basal one third (1/3); four medial veins present, bM3 and dM3 straight; anal cell closed before wing margin. Midtibiae with 1 spur. Etymology. From “ellip- ”, which means elliptic and genus Ragio, referring to the elliptic wings. Gender: masculine. Remarks. Among the snipe flies from the Daohugou locality, the new genus is most similar to Trichorhagio 154 · Zootaxa 4691 (2) © 2019 Magnolia Press Han et al. Zhang, 2013 in appearance, especially in venation and body configuration. But they can be distinguished by elliptic wing (vs. triangular wing in Trichorhagio), and the equal length of bM3 and dM3 (vs. the bM3 longer than dM3). The similar long mouthparts and elliptic wing also present in two other genera Protorhagio Rohdendorf, 1938 and Palaeoarthroteles Kovalev & Mostovski, 1997, which have also been found from the Daohugou. In Protorhagio, costal section between Sc-R1 is obviously longer than that between R1-R2+3, while not distinct in Elliprhagio. Elliprhagio is distinguished from Palaeoarthroteles (Kovalev & Mostovski 1997; Zhang J. 2011) by the configuration of anal cell being closed (vs. open in Palaeoarthroteles); up-curved R2+3 at the middle (vs. almost straight R2+3); straight bM3 and dM3 (vs. curved bM3 and S-shaped dM3 in Palaeoarthroteles); costal section of R1-R2+3 slightly shorter than Sc-R1 (vs. costal section of R1-R2+3 no shorter than Sc-R1 in Palaeoarthroteles) and mesotibiae with 1 apical spur (vs. 2 spurs in Palaeoarthroteles). Moreover, the genus Sinorhagio Zhang, Yang & Ren, 2006, and some species of Palaeobolbomyia (Kovalev 1982; Zhang 2010) in the same locality also possess elliptical wings resembling Elliprhagio. But Sinorhagio can be separated by their straight R2+3, long R4 and R5 branches, and long and narrow d cell. Although Palaeobolbomyia resembles the new genus with similar R2+3, they can be separated by the absence of M3 in the former. Comparing with genera from other localities, the genus Orsobrachyceron Ren, 1998 from Liaoning (China) also has similar long proboscis and venation as those of the Elliprhagio gen. nov., but differs from Elliprhagio in the M3 and M4 strongly converged to a point at the margin of wing and cell cu closed without a short petiole apically. The other two genera Palaeobrachyceron Kovalev, 1981 and Jurabrachyceron Kovalev, 1981 from the Transbaikalia (Upper Jurassic to Lower Cretaceous) show some similarities in the elliptic wings and configurations of venation with Elliprhagio. However, they can be separated by their configuration of M3 and M4: bM3 much shorter than dM3, and M3 parallel to M4. In addition, Palaeobrachyceron has an extremely long and straight R5 that is distinctly different from the new genus. Elliprhagio macrosiphonius sp. nov. (Figs 1, 2, 3) urn:lsid:zoobank.org:act:0807A694-1CDE-4F30-9F72-13FD70E625D5 Holotype. CNU-DIP-NN2015101, well preserved, female. Paratypes. CNU-DIP-NN2015103, antennae and genitalia absent, sex unknown. CNU-DIP-NN2015105, antennae and genitalia absent, sex unknown. CNU-DIPNN2015106, genitalia absent, sex unknown. All the type fossils are housed in the Capital Normal University, Beijing, China. Type locality and horizon. Jiulongshan Formation, in the village of Daohugou, Ningcheng, Inner Mongolia, China (Middle Jurassic). Diagnosis. As for the genus. Description. Moderate-sized flies. Body and legs dark, abdomen covered with short hairs, and legs covered with short bristle. Head: Moderately large, spherical, occiput weakly convex and with short hairs in the lower part. Eyes bare, dichoptic in female (Figs 1A, 2D). The antenna almost as long as the head, scape short, triangular, pedicel longer than scape, bearing short hairs. Flagellum with 10 subsegments, the first segment enlarged obviously, almost two times wider than long. The following three segments slightly narrower than the previous segment in sequence, 5th to 10th segments getting narrower and shorter gradually, the terminal one conical, rather small and short (Figs 1A, B, 2A). Proboscis apparently long, exceeding the length of the head. Labrum long and strongly sclerized, the details of piercing structures indiscernible. Maxillary palpi two-segmented, much shorter than the proboscis. Labium fleshy, labella inflated and enwrapping the distal of the labrum and piercing structures (Figs 1B, 2D). Legs: Coxae of legs bear moderately long setae. All trochanters bearing very short setae. Femora completely covered with short dense setae. Tibiae of legs slightly longer than corresponding femora, covered with short dense setae. Tibial spurs formula 0:1:1 (Fig. 2B, C). Wings: Broad and elliptic, pterostigma blurry, and jugal region undeveloped. Sc ended at the middle of costal vein, crossvein h close to the basal of wing. R1 straight, covered with setae; R2+3 sinuate at the middle, and sharply up-curved distally. Costal section between Sc-R1 nearly as long as that between R1-R2+3. Fork of R4-R5 generally long, and nearly as long as the fork of M1-M2. Crossvein r-m intersecting the upper margin of discal cell at the basal one third (1/3). Four medial veins present. M1 slightly arched. Discal cell narrow, presenting at the middle of the wing. M3 and M4 converged distally. dM3 with equal length to bM3. Cell cu closed and with a very short petiole apiFossil Rhagionidae Zootaxa 4691 (2) © 2019 Magnolia Press · 155 cally (Fig. 2G). Abdomen: Slender, nearly twice as thorax, covered with short setae. Cerci two-segmented, with short setae, 1th segment nearly lobe-like and 2th segment sub-oblong (Figs 1C, 2F). Dimensions. Length of body 6.93‒9.38 mm; head 0.90‒1.08 mm; thorax 1.98‒2.62 mm; abdomen 4.05‒5.68 mm; length of wing 5.89‒6.22 mm; width of wing 2.22‒2.61 mm. Figure 1. Elliprhagio macrosiphonius gen. et sp. nov. Holotype CNU-DIP-NN2015101. A body. B head. C female genitalia. Scale bars: 0.5mm. B, C are under alcohol. Etymology. The specific epithet, “macrosiphonius” is an adjective, and refers to the long mouthparts of this species. Gender: masculine. Remarks. The flagellum of Rhagionidae shows significant morphological diversity that is generally used in the taxonomic treatment. Most extant and Cenozoic rhagionids have little segmented flagellum that the distal part is shrunk to a thin appendage, called arista. However, multi-segmented flagellum is often present in Mesozoic rhagionids. The Middle Jurassic species of Trichorhagio gregarius from Daohugou possesses a nine-segmented flagellum, see figs 1F, 2B in Zhang (2013). It is notable that antennae with 10-segmented flagellum are reported for the first time in E. macrosiphonius gen. et sp. nov. from Daohugou. Therefore, it is deduced that the multi-segmented flagel- 156 · Zootaxa 4691 (2) © 2019 Magnolia Press Han et al. lum is likely a plesiomorphic character of Rhagionidae. Based on these antennal data, it seems that the antennae of rhagionids have a simplified trend leading to fewer and thinner flagellum segments from the Mesozoic to the present, even though phylogenetic relationships among fossil and extant rhagionids have not been clearly elucidated yet. Figure 2. Elliprhagio macrosiphonius gen. et sp. nov. Holotype CNU-DIP-NN2015101. A antenna. B tibial spur of middle leg. C tibial spur of hind leg. D body. E left wing. F female cerci. Scale bars: 1 mm (B–E); 0.5mm (A, F). Discussion As we know, only a few extant Rhagionidae have evolved prolonged proboscis and the mouthpart structure is used for piercing and feeding on blood. Generally, extremely long proboscis within the family is related to the bloodfeeding habit (Kovalev & Mostovski 1997; Lukashevich & Mostovski 2003). Burger (1995) outlined blood-sucking process of extant snipe flies and considered that paired sword-like mandibles for cutting and penetrating, and “retrorse teeth” on outer surface of maxilla should be decisive parts for blood-sucking. Nevertheless, the feeding habit of fossil Rhagionidae was seldom studied because the components of mouthparts in the fossil insects are rarely preserved completely. In 2003, Lukashevich and Mostovski described the species Palaeoarthroteles mesozoicus from the Early Cretaceous and deemed it the most ancient hematophagous brachycerous fly principally derived from the conspicuously long mouthparts and elongate, downcurved palps and unmodified legs (Lukashevich & Mostovski 2003). Interestingly, the new species shows the high similarities of the mouthpart with P. mesozoicus, and it is deduced that E. macrosiphonius, representing the oldest hematophagous rhagionid hitherto, possibly has the similar feeding habit like P. mesozoicus. Almost all fossil snipe flies were established based on the morphological diagnostic characters, especially the wing venation, and this approach and methodology have been widely accepted for the studies of fossil insects (Grimaldi & Cumming 1999). However, the identification is often misguided under the conditions of the limited or Fossil Rhagionidae Zootaxa 4691 (2) © 2019 Magnolia Press · 157 incomplete specimens. Two monotypic genera Trichorhagio and Daohugorhagio were described from Daohugou based on minor differences in appearances, e.g. density of hairs on the body, length of antenna, and features of venation, see figs 1‒4 in Zhang (2013). We have collected a large number of rhagionid fossils from Daohugou, which can be clearly attributed to both genera. After detailed examinations of these specimens, it seems that the diagnostic characters of these two genera emphasized by Zhang (2013) are too equivocal to distinguish them. In the original description, one of the important diagnostic characters is the length of antenna: as long as the head in Trichorhagio vs. half as long as head in Daohugorhagio. In fact, the orientations of head in the specimens of Trichorhagio and Daohugorhagio are distinctly different that could not reflect the actual dimension of the head. Therefore, the length ratio between head and antenna is not a preferred diagnostic character. In addition, the other key character to separate these two genera is configuration of R1, i.e. R1 up-curved near the terminus in Daohugorhagio vs. straight in Trichorhagio. Accordingly, when more specimens were examined, the character of R1 states became continuous, suggesting that R1 has conspicuous intraspecific individual variants. Furthermore, variation of the length and the opening of cell r4 between Daohugorhagio and Trichorhagio is minor and should not be treated as intergeneric variation. The similar variations are also observed in E. macrosiphonius: the opening of cell r4 varies slightly within a narrow range (marked by red arrows in Fig. 3A-D). Based on the afore-mentioned, differences between Trichorhagio and Daohugorhagio should probably just represent interspecific variations, therefore, it is justifiable to treat Daohugorhagio as a synonym to Trichorhagio. The diagnosis of Trichorhagio is emended as follows: body slender, antennae eight or nine-segmented, getting thinner gradually. First flagellomere swollen, thicker than long. R1 straight, costal section of R1-R2+3 much shorter than costal section of Sc-R1. M3 slightly converged to M4. Figure 3. Elliprhagio macrosiphonius gen. et sp. nov., line drawings of wings. A Holotype CNU-DIP-NN2015101. B–D Paratypes. B CNU-DIP-NN2015103. C CNU-DIP-NN2015105. D CNU-DIP-NN2015106. Red arrows mark the opening of cell r4. Scale bars: 1 mm. Acknowledgements We sincerely thank Dr. Kuiyan Zhang for her precious suggestions and help. We are also grateful to Dr. Shih ChungKun for his improvement of our manuscript. And appreciate to the reviewer Dr. M. B. Mostovski and the editor Dr. Vladimir Blagoderov for their critical comments to greatly improve this paper. This research is supported by grants from the National Natural Science Foundation of China (grants 31970383, 31730087, and 41688103), Beijing Natural Science Foundation (grant 5192002), Academy for Multidisciplinary Studies of Capital Normal University, Capacity Building for Sci-Tech Innovation‒Fundamental Scientific Research Funds (grant 19530050144), Program for Changjiang Scholars and Innovative Research Team in University (IRT-17R75), and Support Project of High level Teachers in Beijing Municipal Universities (IDHT20180518). 158 · Zootaxa 4691 (2) © 2019 Magnolia Press Han et al. References Angelini, P.A., Azar, D. & Nel, A. (2016) A new genus and species of snipe fly (Diptera: Rhagionidae) in Lebanese Cretaceous amber. Cretaceous Research, 58, 10–16. https://doi.org/10.1016/j.cretres.2015.09.018 Ansorge, J. (1996) The Upper Liassic insects of Grimmen (Pomerania, north Germany). [Insekten aus dem oberen Lias von Grimmen (Vorpommern, Norddeutschland)]. Neue Paläontologische Abhandlungen, 2, 1–132. Burger, J. (1995) Yellow stone's snipe fly Summer. Yellow stone Science, 3, 2–5. Cockerell, T.D.A. (1908) Two fossil Diptera. The Canadian Entomologist, 40 (6), 173–175. https://doi.org/10.4039/Ent40173-6 Cockerell, T.D.A. (1911) Fossil insects from Florissant, Colorado. Bulletin American Museum of Natural History, 30, 71–82. Cockerell, T.D.A. (1921) Fossil arthropods in the British Museum - VI. Oligocene insects from Gurnet Bay, Isle of Wight. The Annals and Magazine of Natural History, Ninth Series, 7 (42), 453–480. https://doi.org/10.1080/00222932108632550 Evenhuis, N.L. (1994) Catalogue of the Fossil Flies of the world (Inseta: Diptera). Available from: http://hbs.bishopmuseum. org/fossilcat/ (accessed 3 July 2018) Gao, T.P., Shih, C.K., Xu, X., Wang, S. & Ren, D. (2012) Mid-Mesozoic flea-like ectoparasites of feathered or haired vertebrates. Current Biology, 22, 732–735. https://doi.org/10.1016/j.cub.2012.03.012 Grimaldi, D.A. & Cumming, J.M. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1–121. Gu, J.J., Montealeqre-Z, F., Robert, D., Engel, M.S., Qiao, G.X. & Ren, D. (2012) Wing stridulation in a Jurassic katydid (Insecta, Orthoptera) produced low-pitched musical calls to attract females. Proceedings of the National Academy of Sciences, USA, 109 (10), 3868–3873. https://doi.org/10.1073/pnas.1118372109 Handlirsch, A. (1906) Die fossilen insekten und die phylogenie der rezenten Formen. Ein Handbuch für Paläontologen und Zoologen. Engelmann, Leipzig, 640pp. Hong, Y.C., Wang, Z.B. & Sun, W.H. (1992) The stratigraphy and fossil insects of Zhongguan basin, Hebei province. Memoirs of Beijing Natural History Museum, 51, 20–36. Jell, P.A. & Duncan, P.M. (1986) Invertebrates, mainly insects, from the freshwater, Lower Cretaceous, Koonwarra fossil bed (Korumburra Group), South Gippsland, Victoria. In: Jell, P.A. & Roberts J. (Eds.), Plants and invertebrates from the Lower Cretaceous Koonwarra fossil bed, South Gippsland, Victoria. Association of Australian Paleontologists, Sydney, pp. 111–205. Kalugina, N.S. & Kovalev, V.G. (1985) Dipterous insects of Jurassic Siberia. Paleontological Institute, Akademia Nauk, Moscow, pp. 33–133. [in Russian] Kerr, P.H. (2010) Phylogeny and classification of Rhagionidae, with implications for Tabanomorpha Diptera Brachycera. Zootaxa, 2592 (1), 1–133. http://sci-hub.tw/10.11646/zootaxa.2592.1 Kovalev, V.G. (1981) The oldest representatives of the Diptera with short antennae from the Jurassic in Siberia. Paleontological Journal, 15 (3), 84–100. Kovalev, V.G. (1982) Some Jurassic Diptera rhagionids (Muscida, Rhagionidae). Paleontological Journal, 16 (3), 87–99. Kovalev, V.G. (1986) Infraorders Bibionomorpha and Asilomorpha. In: Insects in the early Cretaceous ecosystems of western Mongolia. Trudy Sovmestnaya Sovetsko–Mongol’skaya Paleontologicheskaya Ekpeditsiya, 28, 125–154. [in Russian] Kovalev, V.G. & Mostovski, M.B. (1997) A new genus of snipe flies (Diptera, Rhagionidae) from the Mesozoic of eastern Transbaikalia. Paleontological Journal, 31 (5), 523–527. Kraemer, M.M.S., & Nel, A. (2009) First recorded evidence in the fossil record of snipe flies (Diptera: Rhagionidae) in Cretaceous amber, France. Cretaceous Research, 30 (6), 1367–1375. https://doi.org/10.1016/j.cretres.2009.08.001 Krzemiński, W. & Ansorge, J. (2005) A new rhagionid fly from the Lower Jurassic of Germany. Polskie Pismo Entomologiczne, 74, 369–372. Krzemiński, W. & Krzemińska, E. (2003) Triassic Diptera: descriptions, revisions and phylogenetic relations. Acta Zoologica Cracoviensia, 46 (Supplement), 153–184. Latreille, P.A. (1802) Histoire naturelle, générale et particulière, des crustacés et des insectes. Tome troisième. Familles naturelles des genres. F. Dufart, Paris, 467 pp. Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, o cum characteribus, differentiis, synonymis, locis. Editio Decima, reformata. Laurentii Salvii, Holmiae, 824 pp. Lukashevich, E.D. & Mostovski, M.B. (2003) Hematophagous insects in the fossil record. Paleontological Journal, 37 (2), 153–161. Melander, A.L. (1949) A report on some Miocene Diptera from Florissant, Colorado. American Museum Novitates, 1407, 1– 61. Meunier, F. (1899) Révision des diptères fossiles types de Loew conservés au Musée Provincial de Koenigsberg. Miscellanea Fossil Rhagionidae Zootaxa 4691 (2) © 2019 Magnolia Press · 159 Entomologica, 7, 169–182. Meunier, F. (1902) Études de quelques diptères de l'ambre. Annales des Sciences Naturelles, Zoologie et Paléontologie, 16, 395–404. Meunier, F. (1910) Monographie der Leptiden und der Phoriden des Bernsteins. Jahrbuch der Königlich Preussischen Geologischen Landesanstalt, 30 (2), 64–90. Meunier, F. (1916) Sur quelques diptères (Bombylidae, Leptidae, Dolichopodidae, Conopidae et Chironomidae) de l'ambre de la Baltique.Tijdschrift voor Entomologie, 59, 274–286. Mostovski, M.B. (2000) Contributions to the study of fossil snipe-flies (Diptera: Rhagionidae): The genus Palaeobolbomyia. Paleontological Journal, 34 (Supplement 3), S360–S366. Mostovski, M.B. (2008) Contributions to the study of fossil snipe flies (Diptera: Rhagionidae): The genus Protorhagio. Paleontological Journal, 42 (1), 75–80. Mostovski, M.B. & Jarzembowski, E.A. (2000) The first brachycerous flies (Diptera: Rhagionidae) from the Lower Jurassic of Gondwana. Paleontological Journal, 34 (Supplement 3), S367–S369. Mostovski, M.B., Jarzembowski, E.A., Coram, R.A. & Ansorge, J. (2000) Curious snipe-flies (Dipera: Rhagionidae) from the Purbeck of Dorset, the Wealden of the weald and the Lower Cretaceous of Spain and Transbaikalia. Proceedings of the Geologists’ Association, 111, 153–160. https://doi.org/10.1016/S0016-7878(00)80005-X Nel, A., Perreau, Z. & Doitteau, G. (2016) The oldest representative of the modern snipe fly genus Symphoromyia (Diptera: Rhagionidae). Zootaxa, 4196 (1), 144–150. https://doi.org/10.11646/zootaxa.4196.1.9 Ren, D. (1998) Late Jurassic Brachycera from Northeastern China. Acta Zootaxonomica Sinica, 23, 65–83. Rohdendorf, B.B. (1938) Dipterous insects of the Mesozoic of Karatau. I. Brachycera and part of the Nematocera. Trudy Paleontologicheskogo Instituta Akademii nauk SSSR, 7 (3), 29–67. Rohdendorf, B.B. (1964) The historical development of Diptera. Trudy Paleontologicheskogo Instituta Akademii nauk SSSR, 100, 1–311. Statz, G. (1940) Neue dipteren (Brachycera et Cyclorhapha) aus dem Oberoligozän von Rott. Palaeontographica Abteilung A, 91, 120–174. Théobald, N. (1937) Notes complémentaire sur les insectes fossiles Oligocènes des gypses d'Aix-en-Provence. Bulletin Mensuel de la Société des Sciences de Nancy, 6, 157–178. Wang, Y.J., Labandeira, C.C, Shih, C.K., Ding, Q.L., Wang, C., Zhao, Y.Y. & Ren, D. (2012) An extraordinary case of Jurassic mimicry between a hangingfly and a ginkgo from China. Proceedings of the National Academy of Sciences, USA, 109 (50), 20514–20519. https://doi.org/10.1073/pnas.1205517109 Wootton, R.J. & Ennos, A.R. (1989) The implications of function on the origin and homologies of the dipterous wing. Systematic Entomology, 14, 507–520. https://doi.org/10.1111/j.1365-3113.1989.tb00300.x Yao, Y.Z., Cai, W.Z., Xu, X., Shih, C.K., Engel, M.S., Zhang, X.T., Zhao, Y.Y. & Ren, D. (2014) Blood-feeding true bugs in the Early Cretaceous. Current Biology, 24 (15), 1786–1792. https://doi.org/10.1016/j.cub.2014.06.045 Zaitzev, V.F. (1986) New species of Cretaceous fossil bee flies and a review of paleontological data on Bombyliidae (Diptera). Entomologicheskoe Obozrenie, 65, 815–825. [in Russian] Zetterstedt, J.W. (1842) Diptera Scandinaviæ disposita et descripta. Vol. 1. Officina Lundbergiana, Lundae, 894 pp. Zhang, J.F. (2010) New species of Palaeobolbomyia Kovalev and Ussatchovia Kovalev (Diptera, Brachycera, Rhagionidae) from the Callovian-Oxfordian (Jurassic) Daohugou biota of China: Biostratigraphic and paleoecologic implications. Geobios, 43, 663–669. https://doi.org/10.1016/j.geobios.2010.06.004 Zhang, J.F. (2011) Two new species of Palaeoarthroteles Kovalev and Mostovski (Diptera, Rhagionidae) from the CallovianOxfordian (Jurassic) Daohugou biota of China. Geobios, 44, 635–639. https://doi.org/10.1016/j.geobios.2011.01.006 Zhang, J.F. (2013) Snipe flies (Diptera: Rhagionidae) from the Daohugou Formation (Jurassic), Inner Mongolia, and the systematic position of related records in China. Palaeontology, 56, 217–228. https://doi.org/10.1111/j.1475-4983.2012.01192.x Zhang, J.F. & Li, H.J. (2012) New taxa of snipe flies (Diptera: Brachycera: Rhagionidae) in the Daohugou biota, China. Paleontological Journal, 46, 157–163. Zhang, J.F., Zhang, S. & Li, L.Y. (1993) Mesozoic gadflies (Insecta: Diptera). Acta Palaeontologica Sinica, 32, 662–672. Zhang, K.Y., Yang, D. & Ren, D. (2006) The first snipe fly (Diptera: Rhagionidae) from the Middle Jurassic of Inner Mongolia, China. Zootaxa, 1134 (1), 51–57. https://doi.org/10.11646/zootaxa.1134.1.3 Zhang, K.Y., Yang, D. & Ren, D. (2008) A new genus and species of the Middle Jurassic rhagionids from China (Diptera: Rhagionidae). Biologia, 63 (1), 113–116. https://doi.org/10.2478/s11756-008-0012-4 160 · Zootaxa 4691 (2) © 2019 Magnolia Press Han et al.