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First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae)
Accepted Manuscript First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae) Yuming Liu, Chenyang Cai, Diying Huang PII:
S0195-6671(16)30176-8
DOI:
10.1016/j.cretres.2016.10.007
Reference:
YCRES 3465
To appear in:
Cretaceous Research
Received Date: 24 August 2016 Accepted Date: 16 October 2016
Please cite this article as: Liu, Y., Cai, C., Huang, D., First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae), Cretaceous Research (2016), doi: 10.1016/j.cretres.2016.10.007. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
ACCEPTED MANUSCRIPT
First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae)
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Yuming Liu a, Chenyang Cai b, Diying Huang c, *
a
School of Lisiguang, China University of Geosciences, Wuhan 430074, China
b
Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and
c
SC
Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and
*
M AN U
Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
Corresponding author. Nanjing Institute of Geology and Palaeontology, Chinese Academy of
Sciences, 39 East Beijing Road, Nanjing 210008, China. E-mail address: [email protected] (D.
Abstract
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Huang)
Stratiomyidae, or soldier flies, are a relatively primitive and diverse group of Brachycera, but
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definitive stratiomyids from the Mesozoic are rare. Here we describe the first soldier fly, Lysistrata burmensis sp. nov., in the mid-Cretaceous amber from northern Myanmar. The new species mostly
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resembles Lysistrata emerita Grimaldi and Arillo from the upper Lower Cretaceous Spanish amber and Montsecia martinezdelclosi Mostovski from the Lower Cretaceous Montsec Formation of Spain based on the general characters of venation, antennae and cerci. The new form displays an unusual feature (tibial spurs formula 0-1-0) that is confined exclusively to the modern genus Allognosta Meigen.
Keywords:
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Stratiomyidae Cenomanian Taxonomy
1.
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Myanmar
Introduction
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With nearly 100,000 extant species, Brachycera is an extremely successful group of Diptera that originated in the Triassic. Stratiomyomorpha is one of the most basal groups of Brachycera,
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including the small family Xylomyidae, and the large family Stratiomyidae (Grimaldi and Engel, 2005). Stratiomyidae include over 3000 described species and 82 genera grouped into 12 subfamilies. Adults of solder flies feed mainly on nectar, while most larvae are necrophagous (Yang, 2014). Mesozoic stratiomyid fossils are very sparse. The earliest known soldier fly, Montsecia martinezdelclosi Mostovski, is from the Lower Cretaceous Montsec Formation of Spain (Mostovski,
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1999). In addition, two problematic stratiomyid fossils are described or illustrated from the Lower Cretaceous Yixian Formation of northeastern China (Huang and Lin, 2007) and the Upper Cretaceous of Russian Far East (Blagoderov et al., 2002), but the former considered not belonging to
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Stratiomyidae (Grimaldi and Arillo, 2011). Three Mesozoic representatives are represented in ambers, including Lysistrata emerita Grimaldi and Arillo from the upper Lower Cretaceous amber in the
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Escucha Formation of Alava, Spain (Grimaldi et al., 2011), Cretaceogaster pygmaeus Teskey from the Upper Cretaceous Canadian amber (Teskey, 1971), and an unnamed form from the Upper Cretaceous New Jersey amber (Grimaldi and Cumming, 1999). Besides, multiple stratiomyid larvae are also recorded from the Montsec Formation of Spain (Whalley and Jarzembowski, 1985). Compared the Mesozoic stratiomyid, more fossils have been reported in Cenozoic strata, from the upper Eocene Florissant deposits in Colorado (James 1937; Melander 1949; Carpenter, 1992), mid-Miocene Mexican Chiapas amber (James, 1971; Coty and Nel, 2013), Eocene Baltic amber and
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lower Miocene Dominican amber. (Evenhuis, 1994). Here we describe a new species belonging to Stratiomyidae based on a well-preserved individual from the mid-Cretaceous amber of Myanmar, which represents the first fossil record of
2.
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Stratiomyidae from the deposits.
Material and methods
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The female and male individuals were obtained from the amber deposit in the Hukawng Valley of northern Myanmar. Its geological settings and the amber deposits were summarized by Grimaldi et al.
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(2002) and Ross et al. (2010). Recent U-Pb zircon dating constrained the Burmese amber to a maximum age of 98.79 ± 0.62 Ma (Shi et al., 2012), which indicates a late Albian to early Cenomanian age for the fossilized inclusions. The specimens (NIGP164040 and NIGP164891) are preserved in pieces of relatively clear, yellow amber. The amber containing the holotype is 1.6 cm long, 0.9 cm wide and 0.5 cm thick. The holotype is nearly complete except the right hind leg and
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apex of left wing damaged. The amber containing the allotype is relatively poorly-preserved, with bubbles surround the insects. The ambers were prepared using a mini table-saw, polished with emery papers with different grain sizes, and finally buffed with polishing powder. Photographs were taken
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using a Zeiss Discovery V20 stereo microscope and a Zeiss Axio Imager 2 compound microscope with a digital camera attached, respectively. We follow the nomenclature of (McAlpine, 1981).
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All taxonomic acts established in the present work have been registered in ZooBank (see below), together with the electronic publication under: urn:lsid:zoobank.org:pub:EEBDE07E-A60A-49F9-B091-E1ADC09FFD61.
3.
Systematic palaeontology
Order: Diptera Linné, 1758
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Family: Stratiomyidae Latreille, 1802 Genus: Lysistrata Grimaldi and Arillo, 2011
Lysistrata burmensis sp. nov.
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(urn:lsid:zoobank.org:act:43EB638B-8F28-4D91-B546-122BC3B2047E)
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Type species: Lysistrata emerita Grimaldi and Arillo, 2011
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(Figs. 1–4)
Etymology. The specific epithet is derived from “Burma”, from where the holotype was discovered.
Material. Holotype, NIGP164040; allotype, NIGP164891; both deposited in the Nanjing Institute of
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Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China.
Locality and horizon. Upper Albian to lower Cenomanian, Hukawng Valley, northern Myanmar
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Diagnosis. Antenna with 8 tapering flagellomeres, first flagellomere large, 8th flagellomere elongate; pro- and metatibia without spurs, mesotibia armed with one apical spur; scutellum without spines; R4
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relatively short, forming distinct angle with R5; cell d not very elongate, approximately twice as long as wide; M3 absent; and abdomen elongate, 7-segmented.
Description. Female, slender (Fig. 1), body length 5.08 mm, pilose. Head nearly as wide as thorax (Fig. 2A). Eye bare, big and broadly separated. Ocellar tubercle weakly convex; ocellus not clearly visible. Antenna ca (Fig. 2B). 0.53 mm long; scape and pedicel short, scape 0.03 mm long, pedicel 0.06 mm long, with densely hairs; flagellum 0.40 mm long, first
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flagellomere 4 times length of mid flagellomeres, flagellomeres 2–7 short, nearly in the same size, terminal flagellomere slender, cylinder-shaped, approximately twice as long as mid flagellomeres. Mouthparts well-developed; labellum very broad, with numerous long setae. Occiput flat, pilose.
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Thorax longer than it is wide, densely pilose. Leg slender, fore leg longer than mid- and hind legs pro- and metatibia lacking spur(s), mesotibia with single short apical spur (Fig. 2F); length ratio of tarsomeres 2–5 nearly 9: 6: 4: 6 (Figs. 2D, 2E, 2F); claw simple. Right wing 2.01 mm long and 1.09
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mm wide, covered with microtrichia (Fig. 2C); all veins dark; Sc straight and complete; R4+5
branches from Rs in the apical forth of R2+3, R4+5 separating early, slightly beyond level of discal cell,
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R4 relatively short and form distinct angle with R5, R5 slightly curved backward; M3 absent; CuA2 arcuate, with angle formed by junction of veins CuA2 and A1. Scutellum without spines. Abdomen elongate, covered with bristles and punctations. length 3.60 mm, first abdominal segment very large, 1.43 mm long, length of abdominal segments shortened from base to apex. Cerci long and slender (Fig. 2G), 2-segmented, densely setose.
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Male, body 3.4 mm long, much short than female mainly because of the shape of abdomen (length of head + thorax are very close). Scutellum without spines, with numerous bristles. Venation same as that of female. Abdomen as wide as thorax, with seven segments, display a distinct
Discussion
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4.
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constriction between thorax and abdomen. Genitalia projecting apically, not clearly visible.
The new species closely resembles the extinct species Lysistrata emerita in having similar antennae, wing venation, and elongate and tapered abdomen. In particular, they share similar antennal morphology, including hairy scape and pedicel, prominent basal flagellomere, long and thin terminal flagellomere. L. burmensis bear eight flagellomeres, a character usually found in some extant basal groups (Jorgensen and James, 1968). Instead of eight, L. emerita possess seven flagellomeres, but it cannot be regarded as a key diagnostic character for distinguishing L. burmensis from L. emerita,
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owing to the fact that the antennae of latter are insufficiently preserved. According to Grimaldi and Arillo (2011), the abdomen of L. emerita is elongate, tapered, and with 9 or 10 segments, but seems unclear. Female L. burmensis also possess elongate and tapered abdomen, but with perhaps 7
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abdominal segments, similar to some extant basal subfamilies, including Beridinae, Chiromyzinae and Pachygastrinae. The general vein features are shared by both species, while wings of L.
burmensis are wider than those of another species, which may be caused by different observational
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angles. The most remarkable difference is that the R4 of L. emerita is subparallel to the R5, and the end of R4 is close to the R5. By contrast, in L. burmensis, the R4 forms a distinct angle with the R5,
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and the level of R4 ends in the middle of the R2+3 and R5. In L. emerita, the R2+3 arises from the middle between the base of Rs and r-m, but it arises from the anterior third in L. burmensis. In addition, the cerci of L. burmensis is distinctly longer. Another important difference between them is that the tibial spur formula of L. burmensis is 0-1-0, while it is 0-2-2 in L. emerita; the latter character state is commonly occurring in the majority of modern Stratiomyidae. The 0-1-0 tibial spur
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formula is rare among all extant stratiomyids, and it is usually uniform with a genus (e.g., Allognosta), confined to Archistratiomys and Benhamyia there are all species with one spur (Woodly, 1995). There is no doubt that Lysistrata is closest to some genera of the extant subfamily Beridinae
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by their antennal morphology, number of abdominal segments, and the absence of M3. Lysistratais easily separated from all genera of Beridinae except Allognosta by the spineless scutellum.
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Compared with all modern stratiomyids, the R4 of Lysistrata is relatively long, which probably represents a plesiomorphic character. Therefore, Lysistrata is closest to members of Beridinae, but it cannot be placed in it.
Lysistrata is very similar to Montsecia Mostovski, an extinct genus originally placed in Beridinae and was suggested to be close to the extant Allognosta (Mostovski, 1999). According to Grimaldi (2011), the placement of Montsecia in Beridinae is problematic. Lysistrata share with Montsecia the general antennal morphology (although segmentation remains unclear), the spineless
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scutellum, and the 2-segmented cerci with the second segment more elongate than the first. They also share similar wing venation, but the venation of Montsecia is closer to that of Lysistrata. In addition, the abdomen of Lysistrata is tubular, elongate and gradually tapered posteriorly, distinctly differing
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from the broad and flat abdomen of Montsecia. The tibial spurs of Montsecia remain unknown, so the relationship between Montsecia and Lysistrata requires further studies.
Moreover, Lysistrata also share some similarities (e.g., antennal morphology and spineless
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scutellum) with an undescribed species from the New Jersey amber (Grimaldi and Cumming, 1999). They also have similar wing venation, such as the level of R2+3 and R4 branch. More importantly,
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both genera bear one mesotibial spur. Unlike Lysistrata, the wing of the New Jersey species bears M3, the R4 is slightly waved and branch off latter, the meeting point of CuA2 and A is very close to the wing margin. Besides, the CuP is present in the New Jersey species, and it is absent in all other Mesozoic stratiomyids.
Lysistrata are undoubtedly a primitive lineage of Stratiomyidae, and it may be closely related to
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the extant Allognosta, or possibly to Montsecia from the New Jersey amber. Woodley (1986) suggested that Pachygastrinae is the most basal subfamily, a group with the R4 braches early and nearly parallel to the R5 (also occur in the L. emerita), and the CuA2 is almost perpendicular to the A.
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Woodley (1986) also consider Cretaceogaster pygmaeus as a primitive species, but R4 branches late and the meeting point of CuA2 and A is very close to the wing margin indicate it is a relatively
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advanced species.
5. Concluding remarks
The discovery of the first stratiomyids from Burmese amber increased the distribution and diversity of Mesozoic solder flies. The new species indicates a distinct morphological resemblance to Lysistrata emerita from Alava, Spain, and thus reveals similar palaeoenvironment between these two localities in the mid-Cretaceous. The new species displays more anatomical characters of Lysistrata,
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suggesting a close relationship to recent genus Allognosta and providing important evidence for understanding the evolution of the subfamily Beridinae.
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Acknowledgements We are grateful to two anonymous reviewers for their helpful comments on the manuscript. The work was supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB18030501) and the National Natural Science Foundation of China
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(91514302).
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References
Blagoderov, V.A., Lukashvich, E.D., Mostovski, M.B., 2002. Diptera. In: Rasnitsyn, A.P., Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publishers, Dordrecht, pp. 234–236. Carpenter, F.M., 1992. Superclass Hexapoda. In: Moore, R.C., Kaesler, R.L. (Eds.), Treatise on Invertebrate Paleontology (R) Arthropoda 4, 3/4. Geological Society of America, Boulder, and
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University of Kansas Press, Lawrence, 655 pp
Coty, D., Nel, A., 2013. New fossil Pachygastrinae from Mexican amber (Diptera: Stratiomyidae). Annales de la Société entomologique de France 49 (3), 267–272.
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Evenhuis, N.L., 1994. Catalogue of the Fossil Flies of the World (Insecta: Diptera). Leiden: Backhuys, 600 pp.
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Grimaldi, D.A., Arillo, A., Cumming, J.M., Hauser, M., 2011. Brachyceran Diptera (Insecta) in Cretaceous ambers, part IV: Significant new Orthorrhaphous taxa. ZooKeys 148, 293–332. Grimaldi, D.A., Cumming, J.M., 1999. Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. American Museum of Natural History, New York, 124pp Grimaldi, D.A., Engel, M.S., 2005. Evolution of the Insects, Cambridge University Press, Cambridge, 755 pp. Grimaldi, D.A., Engel, M.S., Nascimbene, P.C., 2002. Fossiliferous Cretaceous amber from
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Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates 3361, 1–72. Huang, D., Lin, Q., 2007. A new soldier fly (Diptera, Stratiomyidae) from the Lower Cretaceous of
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Liaoning Province, northeast China. Cretaceous Research 28 (2), 317–321. James, M.T., 1937. A preliminary review of certain families of Diptera from the Florissant Miocene beds. Journal of Paleontology 11, 241–245.
California Publications in Entomology 63,71–73.
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James, M.T., 1971. A stratiomyid fly (Diptera) from the amber of Chiapas, Mexico. University of
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Jorgensen, N.M., James, M.T., 1968. Evolution of the antenna in the Stratiomyidae (Diptera). Annals of the entomological society of America 61, 870-878.
Latreille, P.A. 1802. Histoire naturelle, générale et particulière, des crustacés et des insects. 467 pp. Linnaeus, C. 1758. Systema naturae per regna trial naturae, secundum classes, ordines, genera, species, cum character ibus, differentiis, synonymis, locis. Ed. 10, Tomus 1. 823 pp
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McAlpine, J. F., 1981. Morphology and Terminology–Adults. In: McAlpine, J.F., Peterson, B.V., Melander, A.L., 1949. A report on some Miocene Diptera from Florissant, Colorado. American Museum Novitates 1407, 1–63.
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Mostovski, M., 1999. A brief review of brachycerous flies (Diptera, Brachycera) in the Mesozoic, with descriptions of some curious taxa. Proceedings of the First Palaeoentomological
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Conference, Moscow 1998, pp. 103–110. Ross, A.J., Mellish, C., York, P., Crighton, B., 2010. Burmese amber. In: Penney, D. (Ed.), Biodiversity of Fossils in Amber from the Major World Deposits. Siri Scientific Press, Manchester, pp. 208–235. Shewell, G.E., Teskey, H.J., Vockeroth, J.R., Wood, D.M. (coords.): Manual of Nearctic Diptera Volume 1. Agriculture Canada Monograph 27, Ottawa, pp. 9–63. Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q., Li, X. 2012.
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Age constraint on Burmese amber based on U–Pb dating of zircons. Cretaceous research 37, 155–163. Teskey, H.J., 1971. A new soldier fly from Canadian amber (Diptera: Stratiomyidae). The Canadian
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Entomologist 103 (12), 1659–1661. Whalley, P.E.S., Jarzembowski, E.A., 1985. Fossil insects from the lithographic limestone of
Montsech (Late Jurassic-Early Cretaceous), Lerida Province, Spain. Bulletin of the British
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Museum, Natural History. Geology 38 (5), 381–412.
Woodley, N.E, 1986, Parhadrestiinae, a new subfamily for Parhadrestia James and Cretaceogaster
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Teskey (Diptera: Stratiomyidae). Systematic Entomology 11, 377–387.
Woodley, N.E., 1995. The genera of Beridinae (Diptera: Stratiomyidae). Memoirs of the Entomological Society of Washington (USA), Washington, 231 pp. Yang, D., 2014. Stratiomyidae of China. China Agricultural University Press, Beijing, 870 pp (in
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Chinese).
Fig. 1. Lysistrata burmensis sp. nov. (NIGP164040) in mid-Cretaceous amber from northern
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Myanmar. A. left lateral view; B. right lateral view. Scale bars: 1 mm.
Fig. 2. Details of Lysistrata burmensis sp. nov. (NIGP164040) in mid-Cretaceous amber from
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northern Myanmar. A. Enlargement of head, showing compound eyes, antennae, and labellum. B. Enlargement of A, showing details of antenna. C. Venation of right forewing. D. Details of protarsi. E. Details of metatarsi. F. Details of mesotibial spurs and mesotarsi. G. Terminalia. Scale bars: 200 µm in A, D, E, F, and G, 100 µm in B, 500 µm in C
Fig. 3. Line drawing of Lysistrata burmensis sp. nov., showing general habitus. Scale bar: 1mm.
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Fig. 4. Allotype (male) of Lysistrata burmensis sp. nov. (NIGP164891) in mid-Cretaceous amber from northern Myanmar. A. General habitus. B. Morphology of right forewing. Scale bars: 1mm in
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A, 500 µm in B.
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S0195-6671(16)30176-8
DOI:
10.1016/j.cretres.2016.10.007
Reference:
YCRES 3465
To appear in:
Cretaceous Research
Received Date: 24 August 2016 Accepted Date: 16 October 2016
Please cite this article as: Liu, Y., Cai, C., Huang, D., First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae), Cretaceous Research (2016), doi: 10.1016/j.cretres.2016.10.007. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
ACCEPTED MANUSCRIPT
First soldier fly from the mid-Cretaceous Burmese amber (Diptera, Stratiomyidae)
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Yuming Liu a, Chenyang Cai b, Diying Huang c, *
a
School of Lisiguang, China University of Geosciences, Wuhan 430074, China
b
Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and
c
SC
Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and
*
M AN U
Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
Corresponding author. Nanjing Institute of Geology and Palaeontology, Chinese Academy of
Sciences, 39 East Beijing Road, Nanjing 210008, China. E-mail address: [email protected] (D.
Abstract
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Huang)
Stratiomyidae, or soldier flies, are a relatively primitive and diverse group of Brachycera, but
EP
definitive stratiomyids from the Mesozoic are rare. Here we describe the first soldier fly, Lysistrata burmensis sp. nov., in the mid-Cretaceous amber from northern Myanmar. The new species mostly
AC C
resembles Lysistrata emerita Grimaldi and Arillo from the upper Lower Cretaceous Spanish amber and Montsecia martinezdelclosi Mostovski from the Lower Cretaceous Montsec Formation of Spain based on the general characters of venation, antennae and cerci. The new form displays an unusual feature (tibial spurs formula 0-1-0) that is confined exclusively to the modern genus Allognosta Meigen.
Keywords:
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Stratiomyidae Cenomanian Taxonomy
1.
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Myanmar
Introduction
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With nearly 100,000 extant species, Brachycera is an extremely successful group of Diptera that originated in the Triassic. Stratiomyomorpha is one of the most basal groups of Brachycera,
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including the small family Xylomyidae, and the large family Stratiomyidae (Grimaldi and Engel, 2005). Stratiomyidae include over 3000 described species and 82 genera grouped into 12 subfamilies. Adults of solder flies feed mainly on nectar, while most larvae are necrophagous (Yang, 2014). Mesozoic stratiomyid fossils are very sparse. The earliest known soldier fly, Montsecia martinezdelclosi Mostovski, is from the Lower Cretaceous Montsec Formation of Spain (Mostovski,
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1999). In addition, two problematic stratiomyid fossils are described or illustrated from the Lower Cretaceous Yixian Formation of northeastern China (Huang and Lin, 2007) and the Upper Cretaceous of Russian Far East (Blagoderov et al., 2002), but the former considered not belonging to
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Stratiomyidae (Grimaldi and Arillo, 2011). Three Mesozoic representatives are represented in ambers, including Lysistrata emerita Grimaldi and Arillo from the upper Lower Cretaceous amber in the
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Escucha Formation of Alava, Spain (Grimaldi et al., 2011), Cretaceogaster pygmaeus Teskey from the Upper Cretaceous Canadian amber (Teskey, 1971), and an unnamed form from the Upper Cretaceous New Jersey amber (Grimaldi and Cumming, 1999). Besides, multiple stratiomyid larvae are also recorded from the Montsec Formation of Spain (Whalley and Jarzembowski, 1985). Compared the Mesozoic stratiomyid, more fossils have been reported in Cenozoic strata, from the upper Eocene Florissant deposits in Colorado (James 1937; Melander 1949; Carpenter, 1992), mid-Miocene Mexican Chiapas amber (James, 1971; Coty and Nel, 2013), Eocene Baltic amber and
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lower Miocene Dominican amber. (Evenhuis, 1994). Here we describe a new species belonging to Stratiomyidae based on a well-preserved individual from the mid-Cretaceous amber of Myanmar, which represents the first fossil record of
2.
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Stratiomyidae from the deposits.
Material and methods
SC
The female and male individuals were obtained from the amber deposit in the Hukawng Valley of northern Myanmar. Its geological settings and the amber deposits were summarized by Grimaldi et al.
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(2002) and Ross et al. (2010). Recent U-Pb zircon dating constrained the Burmese amber to a maximum age of 98.79 ± 0.62 Ma (Shi et al., 2012), which indicates a late Albian to early Cenomanian age for the fossilized inclusions. The specimens (NIGP164040 and NIGP164891) are preserved in pieces of relatively clear, yellow amber. The amber containing the holotype is 1.6 cm long, 0.9 cm wide and 0.5 cm thick. The holotype is nearly complete except the right hind leg and
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apex of left wing damaged. The amber containing the allotype is relatively poorly-preserved, with bubbles surround the insects. The ambers were prepared using a mini table-saw, polished with emery papers with different grain sizes, and finally buffed with polishing powder. Photographs were taken
EP
using a Zeiss Discovery V20 stereo microscope and a Zeiss Axio Imager 2 compound microscope with a digital camera attached, respectively. We follow the nomenclature of (McAlpine, 1981).
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All taxonomic acts established in the present work have been registered in ZooBank (see below), together with the electronic publication under: urn:lsid:zoobank.org:pub:EEBDE07E-A60A-49F9-B091-E1ADC09FFD61.
3.
Systematic palaeontology
Order: Diptera Linné, 1758
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Family: Stratiomyidae Latreille, 1802 Genus: Lysistrata Grimaldi and Arillo, 2011
Lysistrata burmensis sp. nov.
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(urn:lsid:zoobank.org:act:43EB638B-8F28-4D91-B546-122BC3B2047E)
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Type species: Lysistrata emerita Grimaldi and Arillo, 2011
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(Figs. 1–4)
Etymology. The specific epithet is derived from “Burma”, from where the holotype was discovered.
Material. Holotype, NIGP164040; allotype, NIGP164891; both deposited in the Nanjing Institute of
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Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China.
Locality and horizon. Upper Albian to lower Cenomanian, Hukawng Valley, northern Myanmar
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Diagnosis. Antenna with 8 tapering flagellomeres, first flagellomere large, 8th flagellomere elongate; pro- and metatibia without spurs, mesotibia armed with one apical spur; scutellum without spines; R4
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relatively short, forming distinct angle with R5; cell d not very elongate, approximately twice as long as wide; M3 absent; and abdomen elongate, 7-segmented.
Description. Female, slender (Fig. 1), body length 5.08 mm, pilose. Head nearly as wide as thorax (Fig. 2A). Eye bare, big and broadly separated. Ocellar tubercle weakly convex; ocellus not clearly visible. Antenna ca (Fig. 2B). 0.53 mm long; scape and pedicel short, scape 0.03 mm long, pedicel 0.06 mm long, with densely hairs; flagellum 0.40 mm long, first
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flagellomere 4 times length of mid flagellomeres, flagellomeres 2–7 short, nearly in the same size, terminal flagellomere slender, cylinder-shaped, approximately twice as long as mid flagellomeres. Mouthparts well-developed; labellum very broad, with numerous long setae. Occiput flat, pilose.
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Thorax longer than it is wide, densely pilose. Leg slender, fore leg longer than mid- and hind legs pro- and metatibia lacking spur(s), mesotibia with single short apical spur (Fig. 2F); length ratio of tarsomeres 2–5 nearly 9: 6: 4: 6 (Figs. 2D, 2E, 2F); claw simple. Right wing 2.01 mm long and 1.09
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mm wide, covered with microtrichia (Fig. 2C); all veins dark; Sc straight and complete; R4+5
branches from Rs in the apical forth of R2+3, R4+5 separating early, slightly beyond level of discal cell,
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R4 relatively short and form distinct angle with R5, R5 slightly curved backward; M3 absent; CuA2 arcuate, with angle formed by junction of veins CuA2 and A1. Scutellum without spines. Abdomen elongate, covered with bristles and punctations. length 3.60 mm, first abdominal segment very large, 1.43 mm long, length of abdominal segments shortened from base to apex. Cerci long and slender (Fig. 2G), 2-segmented, densely setose.
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Male, body 3.4 mm long, much short than female mainly because of the shape of abdomen (length of head + thorax are very close). Scutellum without spines, with numerous bristles. Venation same as that of female. Abdomen as wide as thorax, with seven segments, display a distinct
Discussion
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4.
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constriction between thorax and abdomen. Genitalia projecting apically, not clearly visible.
The new species closely resembles the extinct species Lysistrata emerita in having similar antennae, wing venation, and elongate and tapered abdomen. In particular, they share similar antennal morphology, including hairy scape and pedicel, prominent basal flagellomere, long and thin terminal flagellomere. L. burmensis bear eight flagellomeres, a character usually found in some extant basal groups (Jorgensen and James, 1968). Instead of eight, L. emerita possess seven flagellomeres, but it cannot be regarded as a key diagnostic character for distinguishing L. burmensis from L. emerita,
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owing to the fact that the antennae of latter are insufficiently preserved. According to Grimaldi and Arillo (2011), the abdomen of L. emerita is elongate, tapered, and with 9 or 10 segments, but seems unclear. Female L. burmensis also possess elongate and tapered abdomen, but with perhaps 7
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abdominal segments, similar to some extant basal subfamilies, including Beridinae, Chiromyzinae and Pachygastrinae. The general vein features are shared by both species, while wings of L.
burmensis are wider than those of another species, which may be caused by different observational
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angles. The most remarkable difference is that the R4 of L. emerita is subparallel to the R5, and the end of R4 is close to the R5. By contrast, in L. burmensis, the R4 forms a distinct angle with the R5,
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and the level of R4 ends in the middle of the R2+3 and R5. In L. emerita, the R2+3 arises from the middle between the base of Rs and r-m, but it arises from the anterior third in L. burmensis. In addition, the cerci of L. burmensis is distinctly longer. Another important difference between them is that the tibial spur formula of L. burmensis is 0-1-0, while it is 0-2-2 in L. emerita; the latter character state is commonly occurring in the majority of modern Stratiomyidae. The 0-1-0 tibial spur
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formula is rare among all extant stratiomyids, and it is usually uniform with a genus (e.g., Allognosta), confined to Archistratiomys and Benhamyia there are all species with one spur (Woodly, 1995). There is no doubt that Lysistrata is closest to some genera of the extant subfamily Beridinae
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by their antennal morphology, number of abdominal segments, and the absence of M3. Lysistratais easily separated from all genera of Beridinae except Allognosta by the spineless scutellum.
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Compared with all modern stratiomyids, the R4 of Lysistrata is relatively long, which probably represents a plesiomorphic character. Therefore, Lysistrata is closest to members of Beridinae, but it cannot be placed in it.
Lysistrata is very similar to Montsecia Mostovski, an extinct genus originally placed in Beridinae and was suggested to be close to the extant Allognosta (Mostovski, 1999). According to Grimaldi (2011), the placement of Montsecia in Beridinae is problematic. Lysistrata share with Montsecia the general antennal morphology (although segmentation remains unclear), the spineless
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scutellum, and the 2-segmented cerci with the second segment more elongate than the first. They also share similar wing venation, but the venation of Montsecia is closer to that of Lysistrata. In addition, the abdomen of Lysistrata is tubular, elongate and gradually tapered posteriorly, distinctly differing
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from the broad and flat abdomen of Montsecia. The tibial spurs of Montsecia remain unknown, so the relationship between Montsecia and Lysistrata requires further studies.
Moreover, Lysistrata also share some similarities (e.g., antennal morphology and spineless
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scutellum) with an undescribed species from the New Jersey amber (Grimaldi and Cumming, 1999). They also have similar wing venation, such as the level of R2+3 and R4 branch. More importantly,
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both genera bear one mesotibial spur. Unlike Lysistrata, the wing of the New Jersey species bears M3, the R4 is slightly waved and branch off latter, the meeting point of CuA2 and A is very close to the wing margin. Besides, the CuP is present in the New Jersey species, and it is absent in all other Mesozoic stratiomyids.
Lysistrata are undoubtedly a primitive lineage of Stratiomyidae, and it may be closely related to
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the extant Allognosta, or possibly to Montsecia from the New Jersey amber. Woodley (1986) suggested that Pachygastrinae is the most basal subfamily, a group with the R4 braches early and nearly parallel to the R5 (also occur in the L. emerita), and the CuA2 is almost perpendicular to the A.
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Woodley (1986) also consider Cretaceogaster pygmaeus as a primitive species, but R4 branches late and the meeting point of CuA2 and A is very close to the wing margin indicate it is a relatively
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advanced species.
5. Concluding remarks
The discovery of the first stratiomyids from Burmese amber increased the distribution and diversity of Mesozoic solder flies. The new species indicates a distinct morphological resemblance to Lysistrata emerita from Alava, Spain, and thus reveals similar palaeoenvironment between these two localities in the mid-Cretaceous. The new species displays more anatomical characters of Lysistrata,
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suggesting a close relationship to recent genus Allognosta and providing important evidence for understanding the evolution of the subfamily Beridinae.
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Acknowledgements We are grateful to two anonymous reviewers for their helpful comments on the manuscript. The work was supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB18030501) and the National Natural Science Foundation of China
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(91514302).
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Fig. 1. Lysistrata burmensis sp. nov. (NIGP164040) in mid-Cretaceous amber from northern
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Myanmar. A. left lateral view; B. right lateral view. Scale bars: 1 mm.
Fig. 2. Details of Lysistrata burmensis sp. nov. (NIGP164040) in mid-Cretaceous amber from
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northern Myanmar. A. Enlargement of head, showing compound eyes, antennae, and labellum. B. Enlargement of A, showing details of antenna. C. Venation of right forewing. D. Details of protarsi. E. Details of metatarsi. F. Details of mesotibial spurs and mesotarsi. G. Terminalia. Scale bars: 200 µm in A, D, E, F, and G, 100 µm in B, 500 µm in C
Fig. 3. Line drawing of Lysistrata burmensis sp. nov., showing general habitus. Scale bar: 1mm.
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Fig. 4. Allotype (male) of Lysistrata burmensis sp. nov. (NIGP164891) in mid-Cretaceous amber from northern Myanmar. A. General habitus. B. Morphology of right forewing. Scale bars: 1mm in
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