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Odonata: Zygoptera) from Mid-Cretaceous Burmese Amber Cretaceous Research 73 (2017) 1e13 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Mesomegaloprepidae, a remarkable new damselfly family (Odonata: Zygoptera) from mid-Cretaceous Burmese amber ** * Diying Huang a, , Dany Azar a, b, Chenyang Cai a, Sibelle Maksoud a, b, AndreNel c, , Günter Bechly d a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, People's Republic of China b Lebanese University, Faculty of Sciences II, Department of Biology, Fanar Matn, P.O. Box 26110217, Lebanon c Institut de Systematique, Evolution, Biodiversite, ISYEB e UMR 7205 e CNRS, MNHN, UPMC, EPHE, Museum national d'Histoire naturelle, Sorbonne Universites, 57 rue Cuvier, CP 50, Entomologie, F-75005, Paris, France d Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-70191, Stuttgart, Germany article info abstract Article history: Mesomegaloprepus magnificus gen. et sp. nov. (Odonata: Zygoptera) is described from more than 14 Received 29 November 2016 specimens in eight pieces of mid-Cretaceous (earliest Cenomanian, ca. 99 Ma) amber from Myanmar. Accepted in revised form 11 January 2017 Possible phylogenetic affinities with the Neotropical Latibasaliidae, Thaumatoneuridae, and Pseudos- Available online 12 January 2017 tigmatinae are discussed, and a relationship with Pseudostigmatinae considered as possible, but because of conflicting evidence separate family status as Mesomegaloprepidae fam. nov. is tentatively preferred. Keywords: The remarkable degree of homoplastic conflict in the wing venational similarities indicates that these Pseudostigmatinae represent relatively weak evidence for phylogenetic relationships. The palaeoecology, including sexual Thaumatoneuridae Latibasaliidae dimorphism in wing colouration, of the new taxon is discussed, and the large number of inclusions fi Cenomanian explained with possible breeding behaviour in association with water- lled tree holes (phytotelmata) of Myanmar the amber tree, similar to extant Pseudostigmatinae. The position of all alleged fossil Thaumatoneuridae Phylogeny are discussed and revised: Eothaumatoneura ptychoptera Pongracz, 1935 from the Eocene Geiseltal lo- cality is restored in Thaumatoneuridae. Cretaceous Euarchistigma and Paleogene Eodysagrion are tenta- tively retained as subfamilies Euarchistigmatinae and Eodysagrioninae in Thaumatoneuridae. Paleogene Dysagrioninae and Petrolestinae are removed from Thaumatoneuridae and attributed to a restored family Dysagrionidae, and Paleocene Latibasaliidae is transferred from Amphipterygoidea to Epallagoidea. © 2017 Elsevier Ltd. All rights reserved. 1. Introduction Here, we describe a remarkable new fossil damselfly taxon, Mesomegaloprepus magnificus gen. et sp. nov. (Zygoptera: Meso- Descriptions of fossil damselflies in Cretaceous amber were megaloprepidae fam. nov.), from eight amber pieces with more relatively rare until the recent palaeoentomological studies on the than 14 conspecific inclusions in 99 Ma Burmese amber. This fossil mid-Cretaceous amber from Myanmar, work which has already shows curious similarities with the extant tropical families Thau- resulted in the description of seven species from the families matoneuridae and Pseudostigmatidae, giving us an occasion to Hemiphlebiidae, Perilestidae, Dysagrionidae, Platystictidae, and discuss the position of the fossil taxa currently considered in these Platycnemididae (Poinar et al., 2010; Huang et al., 2015; Zheng two clades (see Supplementary material). et al., 2016a, b, c, 2017), with further descriptions in preparation by the authors. 2. Materials and methods The fossils were examined and measured using an incident light * Corresponding author. stereomicroscope (Olympus SZX9) and a stereomicroscope (Nikon ** Corresponding author. E-mail addresses: [email protected] (D. Huang), [email protected] (D. Azar), SMZ 1500), as well as a Leitz Wetzlar binocular microscope. Pho- [email protected] (A. Nel), [email protected] (G. Bechly). tographs were taken using a Zeiss Discovery V20 microscope http://dx.doi.org/10.1016/j.cretres.2017.01.003 0195-6671/© 2017 Elsevier Ltd. All rights reserved. 2 D. Huang et al. / Cretaceous Research 73 (2017) 1e13 system. Optical instruments were equipped by camera lucida and it shares several wing venational similarities including a unique digital cameras. The raw digital images were processed with focus triadic branching of vein MP. The gender of the name is stacking software, and figure plates prepared with Adobe masculine. Photoshop™. Diagnosis. Male ligula with distal segment modified to form a very We follow the wing venation nomenclature of Riek and long single flagellum; wings with extensive brown colour (as pre- Kukalova-Peck (1984), emended by Nel et al. (1993) and Bechly served) pattern and very dense wing venation with a several (1996). The higher classification of fossil and extant Odona- hundreds of cells; secondary antenodal crossveins absent, except toptera, as well as characters for family diagnoses are largely based for three accessory crossveins between C and ScP distal of Ax2; on the phylogenetic system proposed by Bechly (1996, 2003). series of five to seven crossveins in antesubnodal space; nodus in a Several recent works that have addressed the higher phylogeny of very basal position, at about 20% of wing length; more than sixty Zygoptera (Dumont et al., 2010; Davis et al., 2011; Dijkstra et al., postnodal crossveins; postnodal and postsubnodal crossveins not 2013, 2014) have been taken into account. aligned; discoidal cell rectangular and crossed by a single crossvein; distal side MAb of discoidal cell not oblique or with reversed obliquity; subdiscoidal cell traversed by two crossveins; base of RP2 2.1. Burmese amber far distal of subnodus; vein CuP between M þ Cu and AA instead of being between M þ Cu and A; bases of RP3/4 and IR2 (midfork) The specimens are preserved in eight pieces of relatively clear, basally recessed midway between arculus and nodus (instead of yellow Burmese amber. The amber pieces were polished before aligned with subnodus); longitudinal wing veins distally distinctly being examined and photographed. The amber piece with paratype curved to posterior wing margin; CuA long, extending well beyond NIGP 161753 has been included in a glass coffin with Canada Bal- mid wing level, with numerous curved posterior branches and sam as medium for better examination. All amber material was characteristical triadic branching pattern of CuA and apical part of legally acquired in Myanmar from local traders with government MP (but not MA, which is unbranched); no intercalary veins be- registration, and legally exported according to the official regula- tween MP and CuA; pterostigma in apical position, short and tions in Myanmar. rectangular (parallel-sided); pterostigmal brace reduced; no lestine Fossil-bearing has mostly been collected from the Hukawng oblique vein ‘O’; absence of several rows of cells between costal Valley in northern Myanmar (formerly known as Burma). For an margin, RA, and RP1 near wing apex. overview of the amber deposit and its geological setting see, e.g., Zherikhin and Ross (2000), Grimaldi et al. (2002), Cruickshank and Mesomegaloprepus magnificus sp. nov. Ko (2003), and Ross et al. (2010). Radiometric UePb zircon dating Figs. 1e8 (Shi et al., 2012) recently constrained this amber to a minimum age Etymology. The specific epithet is after the wonderful coloured of 98.79 ± 0.62 Ma, which is equivalent to the mid-Cretaceous wings of these damselflies. (earliest Cenomanian). The original habitat of the amber forest is Holotype. Holotype female NIGP 164902, allotype male NIGP still controversial, in fact it has originally been assumed to be a 161753, deposited at Nanjing Institute of Geology and Palae- tropical araucarian forest (Grimaldi et al., 2002; Poinar et al., 2007), ontology, P.R. China. possibly with Dipterocarpaceae as another source for the fossil Paratypes. Female NIGP 161754; male NIGP 164903; male NIGP resin. However, the first detailed report on the macromolecular 164904; NIGP 164950; NIGP 164951, all deposited at Nanjing nature and palaeobotanical affinity of burmite (Dutta et al., 2011), Institute of Geology and Palaeontology; SMNS Bu-231 deposited at based on gas chromatography - mass spectrometry, rejected the State Museum of Natural History Stuttgart, Germany. Araucariaceae and Dipterocarpaceae in favour of Pinaceae as the Type-locality and stratum. Tanai Village, Hukawng Valley, Kachin Burmese amber tree. Grimaldi (2016),afterGrimaldi and Ross (in State, northern Myanmar. burmite, mid-Cretaceous (earliest Cen- press), considered ‘based on the abundant inclusions of leafy omanian), ca. 99 Ma (Shi et al., 2012). shoots’ that it was formed by a conifer, and ‘amber produced Diagnosis. As for the genus, since it is presently monotypic (see possibly by Metasequoia (Taxodiaceae) or a close relative’. Even above), wings dark brown with a hyaline apex, in males with pos- though fossil and extant Pinaceae are generally absent from south terior hyaline patch in distal cubital area, and in females with of the equator and from tropical rainforests in particular, a notable narrow hyaline transverse band near base of RP2 (sexual exception is Pinus krempfii from the rainforests of Vietnam dimorphism). (Brodribb and Feild, 2008). Descriptions The family, genus, and species are registered in Zoobank Holotype female NIGP 164902. A head and thorax with
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