NOTICE: This is the author s version of a work that was accepted for publication in Journal of Paleontology. Changes resulting from the publishing

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1 NOTICE: This is the author s version of a work that was accepted for publication in Journal of Paleontology. Changes resulting from the publishing process, such as peer review, editing, corrections or structural formatting may not be reflected in this document. A definitive version was subsequently published in JOURNAL OF PALEONTOLOGY, VOLUME 86, ISSUE 4 (JULY 2012). DOI: /11-126R.1 1

2 CALOSARGUS TALBRAGARENSIS N. SP.: THE FIRST BRACHYCERAN FLY FROM THE JURASSIC OF AUSTRALIA (DIPTERA: ARCHISARGIDAE) S.K. OBERPRIELER 1 and D. K. YEATES 2 1 Division of Evolution, Ecology and Genetics; Research School of Biology; Australian National University, Acton ACT 0200, Canberra, A.C.T Australia; <stef_oberprieler@hotmail.com> 2 CSIRO Ecosystem Sciences, GPO Box 1700, Canberra, A.C.T. 2601, Australia; <david.yeates@csiro.au> ABSTRACT The first fly fossil discovered from the Upper Jurassic Talbragar Fish Bed in Australia is herein described and illustrated. Based on a single, well preserved wing, Calosargus (Calosargus) talbragarensis Oberprieler and Yeates n. sp. is assigned to the extinct brachyceran family Archisargidae, which has so far only been found in China, Kazakhstan and Mongolia. The genus Calosargus Mostovski currently comprises seven species from the Callovian stage (late Middle Jurassic) of China and three species from the Oxfordian stage (early Upper Jurassic) of Kazakhstan. This oldest brachyceran fossil from Australia and first record of the Archisargidae in the southern hemisphere presents evidence that higher flies were present in this region in the Jurassic, when this group is thought to have commenced its diversification. Wing morphology suggests a relationship between this extinct family and Stratiomyidae. KEY WORDS Brachycera, Archisargoidea, Calosargus, Talbragar Fish Bed, fossil wing. INTRODUCTION 2

3 THE TALBRAGAR Fish Bed near Gulgong, New South Wales, is one of only two insectiferous fossil deposits of Jurassic age in Australia (Fig. 1). It is best known for its fossil fish (Woodward, 1895; Wade, 1941; Bean, 2006) and conifers (Walkom, 1921; White, 1981), but in recent years a substantial number of varied insect fossils have been recovered mainly from its southern end (Beattie, 2007; Beattie and Avery, 2012; Beattie and Nel, 2012). Being of Upper Jurassic age (Kimmeridgian, ± 4.27 Ma; Bean, 2006), the insect fauna of the Talbragar Fish Bed forms an important link between the older Lower Jurassic (Sinemurian Toarcian, Ma) Mintaja (Hill River) fauna in Western Australia (Martin, 2008a) and the younger Lower Cretaceous (Aptian, Ma) Koonwarra fauna in Victoria (Jell and Duncan, 1986). The Talbragar Fish Bed lies at the intersection of a terrestrial and an aquatic ecosystem in southern Gondwana. The stratigraphy suggests a shallow freshwater environment (Dulhunty and Eadie, 1969; Percival, 1979; White, 1981; Beattie and Avery, 2012). The insect fauna is dominated by a single species of Protopsyllidae (Hemiptera), with a smaller proportion of other hemipterans (Auchenorryncha and Heteroptera) also present. Coleoptera are also numerous, comprising mainly Archostemata and Polyphaga. Other orders represented are Odonata, Plecoptera, Orthoptera, Neuroptera and Mecoptera (Beattie and Avery, 2012). Diptera are so far only represented by the single wing described here and by a number of unidentified pupae probably belonging to the lower dipteran family Chaoboridae (P. Cranston, personal commun., 2011). Although over 400 insect fossils have been collected to date, only two have been formally described: a presumed hemipteran, Griphologus lowei (Etheridge and Olliff, 1890), originally interpreted as a cicada (Etheridge and Olliff, 1890) but treated as incertae sedis by Handlirsch (1906), and a dragonfly, Austroprotolindenia jurassica Beattie and Nel (Beattie and Nel, 2012). 3

4 During excavations at the Talbragar Fish Bed in April 2010, the first dipteran wing was discovered. Representing the suborder Brachycera, it is not only the first Jurassic record of this suborder in Australia but also the oldest fossil of higher flies known from Australia. Five other, older fly fossils are known from Australia, all representing the suborder Nematocera: a protorhyphid from the Mintaja site in Western Australia (Martin, 2008b) and four species from the Upper Triassic (Carnian, Ma) Mount Crosby Formation in Queensland, assigned to the families Crosaphididae (originally placed in Hemiptera by Evans, 1971, identified as Diptera by Kovalev, 1983; Krzemiñski and Krzemiñska, 2003), Procramptonomyiidae, Paraxymyiidae (Blagoderov, 1999) and Tillyardipteridae (Lukashevich and Shcherbakov, 1999). The Talbragar fossil is also amongst the oldest Brachycera known from Gondwana, only two older specimens of Rhagionidae having been described from the Lower Jurassic Kota Formation in India (Mostovksi and Jarzembowski, 2000). Furthermore, the Talbragar fossil represents the first record of the extinct family Archisargidae from both Australia and Gondwana. To date, this brachyceran family comprises ten genera and 41 species, including the new species described here, and is otherwise known only from Lower to Upper Jurassic localities in China, Kazakhstan and Mongolia (Table 1). In this paper we describe and illustrate the Talbragar dipteran wing as a new species of Archisargidae and discuss its significance in relation to the phylogeny and diversification of Brachycera. MATERIAL AND METHODS The Talbragar Fish Bed is located in the Central West of New South Wales, approximately 30 km north-east of Gulgong (Percival, 1979), at location 32º 10 S., 149º 48 E. 4

5 (Fig. 1). The outcrop is mostly enclosed within a reserve administered by the National Parks and Wildlife Office at Mudgee, New South Wales. This single specimen was collected by the senior author at the southern site of the Talbragar Fish Bed in April 2010 by hand excavation. It was photographed with a Canon EOS 5D Mark II camera using a BK Plus imaging system (Visionary Digital TM ), and the images were edited using Photoshop CS5. Line drawings were prepared using Photoshop Elements 4.0. Wing venation terminology is based on McAlpine (1981). Abbreviations used are C- costa; Sc - subcosta; R 1 - radius; R anterior branch of the radial sector; R posterior branch of the radial sector; Rs - radial sector; M - medial; CuA - cubitus anterior; CuP - cubitus posterior, A - anal; dm - discal cell; br - first basal cell; bm - second basal cell; cup - cell CuP, r-m - radial to medial cross-vein; m-m - medial to medial cross-vein. SYSTEMATIC PALEONTOLOGY Order DIPTERA Linneaus, 1758 Suborder BRACHYCERA Macquart, 1834 Family ARCHISARGIDAE Rohdendorf, 1962 Archisargidae Rohdendorf, 1962, p. 486; Mostovski, 1996a, b, 1997, p. 73; Nagatomi and Yang, 1998, p. 170 (key to genera). Remarks.-- With the description of this species, the family Archisargidae comprises 41 species placed in 10 genera known only from Jurassic localities (Table 1). The new species extends the distribution of the family from the Northern to the Southern Hemisphere. Historically, the family Archisargidae has been treated as a relative of Stratiomyidae (Rohdendorf, 1962), as a synonym of Eremochaetidae (Evenhuis, 1994) and as a member of the superfamily Nemestrinoidea (Nagatomi and Yang, 1998). Most recently it has been placed 5

6 in the superfamily Archisargoidea, together with Eremochaetidae and Kovalevisargidae (Mostovski, 1997; Zhang, 2011). Genus CALOSARGUS Mostovski, 1997 Calosargus Mostovski, 1997, p.74; Nagatomi and Yang, 1998, p. 165 (diagnosis); Zhang et al., 2007b, p. 2 (key to genera). Type species: Calosargus (Calosargus) tatianae Mostovski, Diagnosis. -- Wing slender; Sc ending far beyond middle of wing; R 1 long; R joining with R 1 before wing margin; R 5 ending at wing apex; CuA 1 arising from cell dm; CuP present, CuA 2 short; cell r2+3 closed, r 1 and r wide open, r 5 small, triangular, dm large, cup narrowly open (after Mostovski, 1997; Nagatomi and Yang, 1998). Remarks.-- The genus currently comprises 11 species (Table 1), divided into two subgenera, Calosargus (Calosargus Mostovksi, 1997) and Calosargus (Pterosargus Mostovksi, 1997), which are distinguished by the former having four posterior cells, vein M 4 absent, M 3 present and arising from apical part of dm and br nearly as wide as bm, whereas the latter subgenus has five posterior cells, vein M 3 present and M 4 arising from the basal part of dm, and M 1 and M 2 often longer (Nagatomi and Yang, 1998, p. 166, 170). Calosargus (Calosargus) talbragarensis new species (Figs. 2, 3) Diagnosis.-- Wing most similar to that of C. (C.) validus Zhang, Yang and Ren, 2007 but differing in R 1 being longer than R 2, R 5 ending only marginally before wing apex and origin of Rs at the middle of wing, not basally as in C. (C.) validus. Description.-- Wing elongate, rather narrow, approximately 6 mm long, 4 mm wide. Veins C, Sc, R 1 thick; Sc long, extending beyond middle of wing; R 1 long, arising from about 6

7 middle of wing, opposite to origin of CuA 2, almost twice as long as free section of M 1+2 ; R arising distally to base of dm, ending at R 1 far before wing margin, opposite to split of R 4 and R 5 ; R 4 and R 5 short, divergent; R 5 ending just posteriorly of wing apex; crossvein r-m located in middle of anterior border of dm; M present; CuA 1 arising from dm; CuA 2 arising from bm; CuP present; bm distinctly longer than dm; br as long as bm; A 1 slightly curved distally; cell cup narrowly open. Etymology.-- The species name refers to the type locality. Types.-- Holotype (wing specimen): Talbragar Fish Bed (Upper Jurassic: Kimmeridgian, 151±4 Ma), Gulgong, New South Wales, Australia. S. Oberprieler coll., April In Australian Museum, Sydney (AM), registration number: F Occurrence.-- Only known from the Talbragar Fish Bed. Remarks.-- Poor preservation of the wing makes some features, especially the posterior margin of the wing base and the apex, difficult to interpret. Nonetheless, the venation agrees well with the diagnostic characters of Calosargus (see diagnosis above), especially with R 2+3 joining R 1 well before the wing margin. Within the genus, C. talbragarensis belongs to the subgenus Calosargus (Calosargus) by having 4 posterior cells, M 3 fused with M 4 and arising from the apical part of dm, M and M distinctly separated at base and br nearly as wide as bm. Despite these character agreements, the Talbragar species differs from all others except C. bellus Zhang, Yang and Ren 2007 in having the origin of Rs located nearly opposite to the origin of CuA 2, not proximally of it. It also has a longer, narrower wing than all other species of Calosargus. Comparison.-- The wing of C. talbragarensis is most similar to that of C. (C.) validus Zhang, Yang and Ren, 2007 in the following respects: R ending at R 1 far before wing margin, M and M similar in origin and length, br marginally longer than bm, dm robust and CuA 1 arising from basal 1/3 of dm. Calosargus talbragarensis is also similar to C. (C.) 7

8 bellus Zhang, Yang and Ren, 2007 and C. (C.) antiquus Zhang, Yang and Ren, 2007 in that R arises slightly distally of the base of dm and ends at R 1 far before the wing margin, and it is similar to C. (C.) tenuicellulatus Zhang, Yang and Ren, 2007 in that R ends at R 1 far before the wing margin, R 5 ends just before the wing apex and cells m1, m2 and m3 are elongate. However, in C. (C.) tenuicellulatus cell dm is much more slender and crossvein r-m is located at the basal third of dm. DISCUSSION Most of our knowledge of Jurassic insect faunas comes from fossil deposits in the northern hemisphere, particularly Kazakhstan, Siberia and China (Grimaldi and Engel, 2005). Our finding is an important glimpse of this time period from the Gondwanan southern hemisphere. The discovery of C. talbragarensis has considerable implications for the distribution of the Archisargidae. Zhang (2010) deemed this family to have been a short-lived group that was relatively abundant in Laurasia, and Zhang, Li, Shih and Ren (2010) suggested that it originated in north-eastern China, from where it spread north-westwards in the Middle Jurassic before becoming extinct in the Upper Jurassic. Calosargus talbragarensis shows that the family was more widespread, occurring as far south as Australia. It is possible that Archisargidae were also present in other Gondwanan continents such as South America, Africa, India and Antarctica. The discovery of Archisargidae in the Talbragar Fish Bed also affects our knowledge of dipteran evolution. It is the first evidence for the presence of Brachycera in Australia during the Jurassic, the time period during which this clade is thought to have commenced its diversification (Mostovski, 1999; Wiegmann et al., 2011). The taxonomic placement of the Archisargidae has long been controversial, with Archosargus Rohdendorf having first been regarded as an extinct member of Stratiomyidae (Rohdendorf, 1938), before being placed in a 8

9 separate family and superfamily. We believe that the affinities of the Archisargoidea lie within the Stratiomyomorpha, with the modern families Stratiomyiidae, Xylophagidae and Pantophthalmidae. Crown-group synapomorphies of this lineage are based on a series of larval characters and, in the adults, on the loss of tibial spurs on the fore legs and the costal vein terminating at or before M 2 along the costal margin of the wing (Woodley, 1989; Yeates and Wiegmann, 1999). We are unable to assess any of these synapomorphies in C. tabralgarensis due to its poor state of preservation. Certainly no archisargid fossil with fore legs preserved appears to have tibial spurs. Wing synapomorphies of the family Stratiomyidae are the costalisation of the radial veins and cell dm being reduced in size. Archisargidae appear to have the costa ending at the apex of the wing, and all exhibit costalisation of the radial veins to some extent. The Stratiomyomorpha are thought to have originated around 175 Ma (Grimaldi and Engel 2005; Wiegmann et al., 2011), which coincides with the age of Archisargidae. Therefore it is unclear whether Archisargidae represent an extinct monophyletic group of flies related to stratiomyids or whether they simply are extinct stem stratiomyids. The characteristic wing venation of the Stratiomyidae, featuring costalised radial veins and an origin of Rs opposite to or distally of the origin of CuA 2 (Colless and McAlpine, 1991), can be easily derived from the more generalised venation of Archisargidae. Thus, this new fossil presents further evidence for a relationship between the Archisargidae and Stratiomyidae, as originally suggested by Rohdendorf (1938). If archisargids are indeed extinct (stem-group) Stratiomyidae or Stratiomyomorpha, C. talbragarensis indicates that Australian Stratiomyomorpha are a Gondwanan faunal element rather than originating from more recent immigrations, such as the faunal interchange between Australia and South-East Asia that begun about 25 Ma (Byrne et al., 2011). As the prospect of finding further dipteran fossils in the Talbragar Fish Bed is high, hopefully further excavations will discover additional, more complete fossils of this fly, allowing a more 9

10 thorough description to be made. It is also possible that new techniques, such as micro-ct scanning, will reveal more of the structure preserved in the holotype or in other specimens than can be currently revealed using optical study techniques. ACKNOWLEDGMENTS We thank R. Beattie (Berry, Australia) for organising the excavations at the Talbragar Fish Bed and for providing access to vital fossil specimens, Dr. R. Oberprieler (CSIRO Ecosystem Sciences) for help with the excavations and advice and guidance throughout this project, Dr. Y.Y. Zhen (Australian Museum, Sydney, Australia) for facilitating the loan of fossils from his museum, Dr. L. Ren (Institute of Zoology, Beijing, China) for providing essential Chinese literature, C. Manchester (CSIRO Ecosystem Sciences) for help in photographing the wing and N. M c Grath (landholder) and M. Sharpe (Head Ranger, N.P.W.S., Mudgee) for access to the site. REFERENCES BEAN, L. B The leptolepid fish Cavenderichthys talbragarensis (Woodward, 1895) from the Talbragar Fish Bed (Late Jurassic) near Gulgong, New South Wales. Records of the Western Australian Museum, 23: BEATTIE, R New insect discoveries at the Upper Jurassic Talbragar fish beds, New South Wales, Australia. African Invertebrates, 48: BEATTIE, R., AND S. AVERY Palaeoecology and palaeoenvironment of the Upper Jurassic Talbragar Fossil Fish Bed, Gulgong, New South Wales, Australia, Alcheringa, in press. BEATTIE, R., AND A. NEL A new dragonfly, Austroprotolindenia jurassica (Odonata: Anisoptera), from the Upper Jurassic of Australia. Alcheringa, DO1:10.180/

11 BLAGODEROV, V. A New Bibionomorpha from the Triassic of Australia and Jurassic of Central Asia with notes on Paraxymyiidae Rohdendorf (Insecta, Diptera). Proceedings of the First Palaeoentomological Conference, Moscow 1998 (ed. P. Vrsanski). AMBA Projects lnternational, Bratislava, p BYRNE, M., D. A. STEANE, L. JOSEPH, D. K. YEATES, G. J. JORDAN, D. CRAYN, K. ALPIN, D. J. CANTRILL, L. G. COOK, M. D. CRISP, J. S. KEOGH, J. MELVILLE, C. MORITZ, N. PORCH, K. SNIDERMAN, P. SUNNUCKS AND P..H. WESTON Decline of a biome: evolution, contraction, fragmentation, extinction and invasion of the Australian mesic zone biota. Journal of Biogeography, 38: COLLESS, D. H., AND D. K. MCALPINE Chapter 39: Diptera In The Insects of Australia, 2nd ed., Vol. 2 (CSIRO, ed.). Melbourne University Press, Carlton, Victoria. DULHUNTY, J. A., AND J. EADIE Geology of the Talbragar fossil fish bed area. Journal and Proceedings of the Royal Society of New South Wales, 102: 1 4. ETHERIDGE, R. J., AND A. S. OLLIFF The Mesozoic and Tertiary Insects of New South Wales. Memoirs of the Geological Society of New South Wales, Palaeontology, 7: EVANS, J. W Some Upper Triassic Hemiptera from Mount Crosby, Queensland. Memoirs of the Queensland Museum, 16: EVENHUIS, N. L Catalogue of the Fossil Flies of the World (Insecta: Diptera). Backhuys Publishers Leiden. Netherlands. GRIMALDI, D. A., AND M. S. ENGEL Evolution of the Insects. Cambridge University Press, New York. HANDLIRSCH, A Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch für Paläontologen und Zoologen. Fasc.1 7. Wilhelm Engelmann, Leipzig, 1430 p. [p published in 1906, p in 1907, p in 1908.] HONG, Y Middle Jurassic Fossil Insects in North China, Geological Publishing House, Beijing. [in Chinese] 11

12 JELL, P. A., AND P. M. DUNCAN Invertebrates, mainly Insects, from the Freshwater, Lower Cretaceous, Koonwarra Fossil Bed (Korumburra Group), South Gippsland, Victoria. In P. A. Jell and J. Roberts (eds), Plants and Invertebrates from the Lower Cretaceous Koonwarra Fossil Bed, South Gippsland, Victoria, p Association of Australian Palaeontologists: Sydney. KOVALEV, V. G A new subfamily of the Diptera from the Triassic deposit of Australia and its presumable descendants (Diptera, Crosaphididae faro. nov., Mycetobiidae). Entomologicheskoe Obozrenie, 62: (In Russian). KRZEMIÑSKI, W. AND E. KRZEMIÑSKA Triassic Diptera: descriptions, revisions and phylogenetic relations. Acta zoologica cracoviensia, (suppl. Fossil Insects), 46: LINNAEUS, C Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Tomus I. Laurentii Salvii, Holmiae, [1-4], p LUKASHEVICH. E. D., AND D. E. SHCHERBAKOV A new Triassic family of Diptera from Australia. Proceedings of the First Palaeoentomological Conference, Moscow 1998 (ed. P. Vrsanski). AMBA Projects International, Bratislava, p MACQUART, P. J. M Histoire naturelle des insectes. Dipteres. Paris, Librairie Encyclopedique de Roret Tome 1, p MARTIN, S. K. 2008a. Hill River rediscovered: Early Jurassic insects of the Perth Basin, Western Australia. Alavesia, 2: MARTIN, S. K. 2008b. A new protorhyphid fly (Insecta: Diptera: Protorhyphidae) from the Lower Jurassic of the Perth Basin, Western Australia. Alavesia, 2: MCALPINE, J Morphology and Terminology - Adults. In J. McAlpine, B. Peterson, G. Shewell, H. Teskey, J. Vockeroth et al. (eds.), Manual of Nearctic Diptera Volume 1, p Ottawa: Research Branch, Agriculture Canada. 12

13 MOSTOVSKI, M. B. 1996a. New species of the genus Mesosolva Hong, 1983 (Diptera, Archisargidae) from the Jurassic of Kazakshtan and Mongolia. In Morales (ed.), The Continental Jurassic. Museum of Northern Arizona Bulletin, 60: MOSTOVSKI, M. B. 1996b. To the knowledge of Archisargoidea (Diptera, Brachycera). Families Eremochaetidae and Archisargidae. Russian Entomological Journal, 5: MOSTOVSKI, M. B To the knowledge of fossil dipterans of the superfamily Archisargoidea (Diptera, Brachycera). Paleontologicheskii Zhurnal, 1: MOSTOVSKI, M. B A brief review of brachycerous flies (Diptera, Brachycera) in the Mesozoic, with descriptions of some curious taxa. Proceedings of the First Palaeoentomological Conference, Moscow 1998 (ed. P. Vrsanski). AMBA Projects lnternational, Bratislava, p MOSTOVSKI, M.B., AND E. A. JARZEMBOWSKI The first brachycerous flies (Diptera: Rhagionidae) from the Lower Jurassic of Gondwana. Paleontological Journal, 34: NAGATOMI, A., AND D. YANG A review of extinct Mesozoic genera and families of Brachycera (Insecta, Diptera, Orthorrhapha). Entomologists Monthly Magazine, 134: NAUMANN, I. D., AND P. B CARNE (eds.) The Insects of Australia, 2nd ed., Vol. 2 (CSIRO, ed.), Melbourne University Press, Carlton, Victoria. PERCIVAL, I. G The geological heritage of New South Wales. Sydney, Australian Heritage Commission and the Planning and Environment Commission of New South Wales, p ROHDENDORF, B. B Dipterous insects of the Mesozoic of Karatau. I. Brachycera and part of the Nematocera. Trudy Paleontologischeskogo Instituta, Akademii Nauk SSSR, 7: (in Russian). ROHDENDORF, B. B Fundamentals of Palaeontology, Volume 9: Arthropoda Tracheata, Chelicerata. Amerind Publishing Company, New Delhi, India (1991 English translation from Osnovy Paleontologii. Tom 9: Chlenistonogie, Trakheinye i Khelitserovye. Akademiya Nauk SSR, Moscow, 1962). WADE, R. T The Jurassic fishes of New South Wales. Journal and Proceedings of the Royal Society of New South Wales, 75:

14 WALKOM, A. B Mesozoic floras of New South Wales. Part 1. Fossil plants from Cockabutta Mountain and Talbragar. Memoirs of the Geological Society of New South Wales, Palaeontology, 12: WIEGMANN, B. M., M. D. TRAUTWEIN, I. S. WINKLER, N. B. BARR, J. KIM, C. LAMBKIN, M. A. BERTONE, B K. CASSEL, K. M. BAYLESS, A. M. HEIMBERG, B. M. WHEELER, K. J. PETERSON, T. PAPE, B. J. SINCLAIR, J. H. SKEVINGTON, V. BLAGODEROV, J. CARAVAS, S. N. KUTTY, U. SCHMIDT-OTT, E. KAMPMEIER, F. C, THOMPSON, D. A. GRIMALDI, A. T. BECKENBACH, G. W. COURTNEY, M. FRIEDRICH, R. MEIER, AND D. K. YEATES Episodic radiations in the fly tree of life. Proceedings of the National Academy of Science of the USA, 108: WHITE, M. E Revision of the Talbragar Fish Bed flora (Jurassic) of New South Wales. Records of the Australian Museum, 33: WOODLEY, N Phylogeny and classification of the Orthorrhaphous Brachycera, p In J. F. McAlpine (ed.), Manual of Nearctic Diptera, Research Branch, Agriculture Monograph No. 32, Vol 3, (Canadian Government Publishing Centre, Hull, Canada). WOODWARD, A. S The fossil fishes of the Talbragar Beds (Jurassic). Memoirs of the Geological Survey of New South Wales, Palaeontology, 9: YEATES, D. K., AND B. M. WIEGMANN Congruence and controversy: Toward a higher-level phylogeny of Diptera. Annual Review of Entomology, 44: ZHANG, J. F Records of bizarre Jurassic brachycerans in the Daohugou biota, China (Diptera, Brachycera, Archisargidae and Rhagionemestriidae). Palaeontology, 53: ZHANG, J. F Three distinct but rare kovalevisargid flies from the Jurassic Daohugou biota, China (Insecta, Diptera, Brachycera, Kovalevisargidae). Palaeontology, 54: ZHANG, J. F., AND H. C. ZHANG Two new species of archisargids (Insecta: Diptera: Archisargidae) from the Upper Jurassic Daohugou Formation (Inner Mongolia, Northeastern China). Paleontological Journal, 37(4):

15 ZHANG, K., T. LI, C. SHIH AND D. REN Chapter 20: Diptera Vampires for Dinosaurs. In Silent Stores - Insect Fossil Treasures from Dinosaur Era of the Northeastern China. Science Press, Beijing, p ZHANG, K., J. LI, D. YANG AND D. REN A new species of Archirhagio Rohdendorf, 1938 from the Middle Jurassic of Inner Mongolia of China (Diptera: Archisargidae). Zootaxa, 1984: ZHANG K., D. YANG AND D. REN Middle Jurassic fossils of the genus Sharasargus from Inner Mongolia, China (Diptera: Archisargidae). Entomological Science, 11: ZHANG, K., D. YANG, D. REN AND C. SHIH. 2007a. The earliest species of the extinct genus Archisargus from China (Diptera: Brachycera: Archisargidae). Annales Zoologici, 57: ZHANG, K., D. YANG, D. REN AND C. SHIH. 2007b. The oldest Calosargus Mostovski, 1997 from the Middle Jurassic of China (Diptera: Brachycera: Archisargidae). Zootaxa, 1645: ZHANG, K., D. YANG, D. REN AND C. SHIH New archisargids from China (Insecta: Diptera). Entomological Science, 13: FIGURE 1 Map of New South Wales showing the general location of the Talbragar Fish Bed (after Beattie and Avery, 2012). 15

16 FIGURE 2 Calosargus (Calosargus) talbragarensis n. sp. (F , AM), wing fossil. Scale bar 1 mm. FIGURE 3 Calosargus (Calosargus) talbragarensis n. sp. (F , AM), line drawing of wing venation, image reversed. Scale bar 2 mm. 16

17 TABLE 1 Genera and species of Archisargidae (type species in bold) with ages and localities. Genus Species Age (Ma) Stage Locality Country Reference Archirhagio obscurus Oxfordian Karatau Kazakhstan Rohdendorf 1938 Archirhagio striatus Callovian Daohuguo village China Zhang and Zhang 2003 Archirhagio zhangi Callovian Daohuguo village China Zhang et al Archisargus maximus Oxfordian Karatau Kazakhstan Mostovski 1997 Archisargus pulcher Oxfordian Karatau Kazakhstan Rohdendorf 1938 Archisargus spurivenius Callovian Daohuguo village China Zhang et al. 2007a Archisargus strigatus Callovian Daohuguo village China Zhang et al. 2007a Brevisolva daohugouensis Callovian Daohuguo village China Zhang Yang Ren and Shih 2010 Calosargus ( Calosargus) niger Oxfordian Karatau Kazakhstan Mostovski 1997 Calosargus (Calosargus) antiquus Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) bellus Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) daohugouensis Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) hani Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) tatianae Oxfordian Karatau Kazakhstan Mostovski 1997 Calosargus (Calosargus) tenuicellulatus Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) validus Callovian Daohuguo village China Zhang et al. 2007b Calosargus (Calosargus) talbragarensis ±4.27 Kimmeridgian Talbragar Fish Bed Australia n. sp. Calosargus (Pterosargus) thanasymus Oxfordian Karatau Kazakhstan Mostovski 1997 Calosargus (Pterosargus) sinicus Callovian Daohuguo village China Zhang 2010 Mesosolva daohugouensis Callovian Daohuguo village China Zhang and Zhang 2003 Mesosolva jurassica Callovian Daohuguo village China Zhang Yang Ren and Shih 2010 Mesosolva longivena Oxfordian Karatau Kazakhstan Mostovski 1996a 17

18 Mesosolva parva Callovian Yujiagou village China Hong 1983 Mesosolva sinensis Callovian Daohuguo village China Zhang Yang Ren and Shih 2010 Mesosolva angustocellulata Oxfordian Karatau Kazakhstan Mostovski 1996b Mesosolva balyshevae Oxfordian Karatau Kazakhstan Mostovski 1996b Mesosolva dolosa Oxfordian Karatau Kazakhstan Mostovski 1996b Mesosolva hennigi Oxfordian Karatau Kazakhstan Mostovski 1996b Mesosolva imperfecta Oxfordian Karatau Kazakhstan Mostovski 1996b Mesosolva rohdendorfi Oxfordian Karatau Kazakhstan Mostovski 1996b Mostovskisargus portentosus Callovian Daohuguo village China Zhang 2010 Mostovskisargus signatus Callovian Daohuguo village China Zhang 2011 Ovisargus gracilis Oxfordian Karatau Kazakhstan Mostovski 1996b Parvisargus malus Oxfordian Karatau Kazakhstan Mostovski 1996b Parvisargus peior Oxfordian Karatau Kazakhstan Mostovski 1996b Prosolva huabiensis Callovian Yujiagou village China Hong 1983 Prosolva karataviensis Oxfordian Karatau Kazakhstan Mostovski 1996b Sharasargus eximius Callovian Daohuguo village China Zhang et al Sharasargus fortis Callovian Daohuguo village China Zhang et al Sharasargus ruptus Tithonian Shar-Teg Mongolia Mostovski 1996b Sharasargus spiniger Oxfordian Karatau Kazakhstan Mostovski 1996b 18