Research 72 (2017) 105e109

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Cretaceous Research

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Short communication The first aeshnoid dragonfly (: Anisoptera: ) from mid-Cretaceous Burmese amber

* ** Daran Zheng a, b, , Su-Chin Chang b, , Edmund A. Jarzembowski a, c, Bo Wang a, d a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China b Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region c Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK d Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1, Beichen West Road, Beijing 100101, China article info abstract

Article history: A new true dragonfly, Cretaeshna lini gen. et sp. nov., is described based on a forewing from mid- Received 25 September 2016 Cretaceous Burmese amber. Cretaeshna is probably a member of Telephlebiidae: Telephlebiinae, but Received in revised form differs from the latter in having a weakly-defined IR1 and a short pterostigma. Cretaeshna lini is the first 26 November 2016 aeshnid dragonfly to be found as an amber inclusion and the third Cretaceous true dragonfly recorded in Accepted in revised form 20 December 2016 amber. Our find augments the diversity of Mesozoic true dragonflies, and enhances our understanding of Available online 26 December 2016 the palaeogeographic distribution of aeshnid dragonflies. © 2016 Elsevier Ltd. All rights reserved. Keywords: Telephlebiidae Anisoptera Cenomanian Cretaceous Burmese amber

1. Introduction 2016a). No other Cretaceous true dragonflies have been previ- ously recorded as the amber inclusions, and only two Cenozoic The world-wide, recent Family is one of the most inclusions have been described: a fragmentary wing from the diverse amongst the true dragonflies, totalling 51 genera and over lowermost Eocene amber of France (Fleck et al., 2000) and a 456 species (von Ellenrieder, 2002; Dijkstra et al., 2013; Walia et al., probably missing specimen from the Eocene of Baltic amber 2016). The fossil Aeshnidae have also been recorded worldwide, (Schadel€ and Bechly, 2016). In this paper, a new true dragonfly, and the oldest record is inferna Pritykina, 1977 from Cretaeshna lini gen et sp. nov., is described from Burmese amber. the Lower Cretaceous of Russia (Pritykina,1977; Bechly et al., 2001), but they were more widely distributed during the Cenozoic (Nel 2. Material and methods et al., 1994; Bechly et al., 2001; Petrulevicius et al., 2010; Prokop et al., 2016). The specimen described herein was collected in the Hukawng Odonatans are comparatively rare as amber inclusions and true Valley of Kachin Province, Myanmar (locality in Kania et al., 2015: fl dragon ies are extremely rare (Zheng et al., 2016a). Only two fig. 1), where the amber was dated at 98.79 ± 0.62 Ma based on fl fragmentary true dragon ies have been recorded from mid- UePb zircon dating from the volcanoclastic matrix (Shi et al., 2012). Cretaceous Burmese amber attributed to the families Damselflies are more common than true dragonflies as Burmese € (Schadel and Bechly, 2016) and Gomphaeschnidae (Zheng et al., amber inclusions, the latter being quite rare. Damselflies in Burmese amber are represented by several extant families, viz., Hemiphlebiidae (Zheng et al., 2016b), (Poinar et al., 2010; Huang et al., 2015; Zheng et al., in press), Platysticti- * Corresponding author. Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region. dae (Huang et al., 2015; Zheng et al., 2016d), and fl ** Corresponding author. (Zheng et al., 2016e). Two extinct dysagrionid damsel ies (Zheng E-mail address: [email protected] (D. Zheng). et al., 2016b,f) and a damsel-dragonfly(Bechly and Poinar, 2013) http://dx.doi.org/10.1016/j.cretres.2016.12.013 0195-6671/© 2016 Elsevier Ltd. All rights reserved. 106 D. Zheng et al. / Cretaceous Research 72 (2017) 105e109 have also been described from Burmese amber. The most common Diagnosis. As for , by monotype. seen species is Burmahemiphlebia zhangi Zheng et al., 2016b, which Description. Wing hyaline (Figs. 1e3), preserved length, 15.3 mm; is a member of the Hemiphlebiidae (Zheng et al., 2016b). width at level of N, 5.3 mm; distance from N to Pt, 11 mm, from The amber piece containing the dragonfly is yellow and trans- Pt to wing apex, 3.2 mm. Pt short (Fig. 3A), less than two cells parent. Photographs were taken using a Zeiss Stereo Discovery V16 long, i.e. 1.3 mm long and 0.4 mm wide, narrowly elongate, with microscope system with Zen software. In most instances, incident basal side somewhat more oblique than distal side. Pterostigmal and transmitted light were used simultaneously. All images are brace quite long, nearly two times length of basal side of Pt, digitally stacked photomicrographic composites of approximately weakly oblique, aligned with basal side of Pt. 11 postnodal 40 individual focal planes obtained using the free software crossveins present before Pt, not aligned with nine postsubnodal Combine ZP for a better illustration of the 3D structures. The line crossveins; three postnodal and three postsubnodal crossveins drawings were prepared from photographs using image-editing present distal of Pt, non-aligned. One oblique crossveins ‘O’ software (CorelDraw X7 and Adobe Photoshop CS6). The spec- (Fig. 3B) present between RP2 and IR2, one cell distal of Sn, i.e. imen is housed in the Nanjing Institute of Geology and Palae- 0.9 mm distally. Base of RP2 slightly distal of Sn, 0.4 mm away. ontology, Chinese Academy of Sciences (NIGPAS). All taxonomic Area between RP2 and IR2 with one row of cells inbetween. IR1 acts established in the present work have been registered in Zoo- short (Fig. 3C),zigzagged,originatingjustbelowbaseofPtbrace, Bank (see below), together with the electronic publication LSID: but forked distally. RP1 and RP2 basally parallel with Pt, with urn:lsid:zoobank.org:pub:7F7B1793-24AF-4CBF-949E- only one row of cells inbetween before Pt, but divergent distally 12509271B3F0. with two rows just below Pt brace, and 11 cells counted along The nomenclature of the dragonflywingvenationusedinthis wing margin. IR2 forked basal of Pt (Fig. 3C) with two row of paper is based on the interpretations of Riek (1976) and Riek and cells between branches below Pt but six rows along posterior Kukalova-Peck (1984),asmodified by Nel et al. (1993) and wingmargin.Rsplwelldefined (Fig. 3C), originating three cells Bechly (1996). Wing abbreviations are as follows: Cr, nodal basal of IR2, parallel with lower branch of IR2 with one row of crossvein; IR, intercalary radial vein; MA, median anterior; Mspl, cells inbetween; RP3/4 and MA running almost parallel basally, median supplement; MP, median posterior; N, nodus; Pt, pter- with one row of cells in between mostly; area between Rspl and ostigma; RA, radius anterior; RP, radius posterior; Rspl, radial RP3/4 with two tows of cells inbetween before IR2 fork, but supplement; ScP, subcosta posterior; Sn, subnodal crossvein. The expanded distally with 14 cells along wing margin. Postdiscoidal phylogeny and classification is based on von Ellenrieder (2002) area with three cells below N and at least 12 cells counted along for the recent genera of Aeshnidae and Bechly (2016) for wing margin. Mspl well defined (Fig. 3D), curved, parallel with higher taxa. MA with one row of cells inbetween; area between Mspl and MP with two rows of cells inbetween below N, but expanded distally with at least 11 cells along wing margin. 3. Systematic palaeontology

Order Odonata Fabricius, 1793 4. Discussion Suborder Anisoptera Selys-Longchamps, 1854 Superfamily Aeshnoidea Leach, 1815 Cretaeshna lini having a well-defined Rspl and Mspl, a basal Family Telephlebiidae Cockerell, 1913 lestine oblique vein shifted basally close to the subnodus, vein IR2 with a distal dichotomous fork, means the wing can be attributed to New genus Cretaeshna gen. nov. the clade Euaeshnodea Bechly et al., 2001 (Brachytronidae (urn:lsid:zoobank.org:act:EC77231A-7CEE-4322-BDA2- Cockerell, 1913 þ Aeshnoidea Leach, 1815). C. lini can be excluded 2017AF53D3A7) from Brachytronidae since the latter have a reduced pterostigmal Type species: Cretaeshna lini sp. nov. brace vein and a basally prolonged pterostigma. Aeshnoidea consist Etymology. A combination of ‘Cretaceous’ and , gender of two families (Bechly, 2016): Telephlebiidae Cockerell, 1913 neuter. (Austroaeschninae Bechly, 1996 þ Telephlebiinae Cockerell, 1913) Diagnosis. Pt well braced, short, covering less than two cells; Pt and Aeshnidae Leach, 1815. Aeshnidae can be excluded from brace well aligned with basal side of Pt; RP2 curved; IR2 forked, consideration since they have a characteristic bulge in the distal with two rows of cells inbetween below Pt base; IR1 short and part of MA in both pairs of wings, distinctly curved Rspl and Mspl zigzagged, not well-defined; Rspl well-defined, parallel with lower with more than one row of cells between them and IR2 or MA, branch of IR2 with one row of cells inbetween; Rspl originating differentiating them from C. lini. three cells basal of IR2 fork; Mspl well-defined, very gently undu- Within Telephlebiidae, the subfamily Telephlebiinae has the lated, parallel with MA with one row of cells inbetween; one row of median space traversed by crossveins and hypertriangles traversed cells between MA and RP3/4 basal of IR2 fork. by at least three crossveins in both pairs of wings, whilst the sub- family Austroaeschninae has no autapomorphies (Bechly, 2016). Cretaeshna lini sp. nov. The absence of the wing base in C. lini makes it difficult to compare (urn:lsid:zoobank.org:act:E0A80A5C-8F6D-4B59-9629- it to the above subfamilies. Compared with Telephlebiidae, C. lini 540AE084D55E) has a forked IR2, pterostigmal brace present and aligned with the Figs. 1e3 base of the pterostigma, MA parallel with RP3/4 to the wing margin, Etymology. The specific name is in honour of Prof. Qibin Lin, a well-defined Mspl nearly parallel to MA, a curved RP2, a well- palaeoentomologist. defined Rspl approximately parallel to IR2, all shared by the Holotype. NIGP164824, a fragmentary forewing with distal half following members of the family following von Ellenrieder (2002): preserved and the posterior margin slightly deformed, gender un- Spinaeschna Theischinger, 1982, Tillyard, 1916, Aus- known; deposited in the Nanjing Institute of Geology and Palae- troaeschna Selys, 1883 and Planaeschna Mclachlan, 1896 in Austro- ontology, Chinese Academy of Sciences, China. aeschninae, Caliaeschna Selys, 1883, Gynacanthaeschna Fraser, 1922, Locality and Horizon. Hukawng Valley, Kachin Province, Myanmar; Selys, 1883, Periaeschna Martin, 1908, Petaliaeschna lowermost Cenomanian, lowermost Upper Cretaceous. Fraser, 1927 and Selys, 1883 in Telephlebiinae. These D. Zheng et al. / Cretaceous Research 72 (2017) 105e109 107

Fig. 1. Cretaeshna lini gen. et sp. nov., holotype, NIGP164824, photograph of specimen.

Fig. 2. Cretaeshna lini gen. et sp. nov., holotype, NIGP164824, line drawing showing venations of forewing. characters are also shared by Antipodophlebia Fraser, 1960 and Within Telephlebiinae, the type genus Telephlebia can be easily Dendroaeschna Tillyard, 1916 after von Ellenrieder (2002), which excluded from Cretaeshna by the presence of brown wing bands are not included in the classifications of Bechly (2016). Anti- and a long pterostigma in the former (Theischinger and Hawking, podophlebia and Dendroaeschna can be easily excluded since they 2006). Caliaeschna and Periaeschna differ from Cretaeshna in hav- have a well-defined IR1, a long Rspl originating more basally, and ing a well-defined IR1, a short Pt brace, a long pterostigma, and two two row of cells between RP3/4 and MA below Rspl (Tillyard, 1916; rows of cells between RP3/4 and MA below Rspl in the former Theischinger, 1977), differing from Cretaeshna. (Wilson and Xu, 2008; Xu, 2007, 2012). Gynacanthaeschna has the Within Austroaeschninae, Notoaeschna and Spinaeschna are Pt brace situated distal to the basal side of the pterostigma (Fraser, unlike Cretaeshna in having a long IR1, more than two row of cells 1936; Wilson and Xu, 2008) showing a difference between it and between the two branches of IR2 just below the pterostigma and Cretaeshna. Cephalaeschna differs from Cretaeshna in having a long Rspl originating more basally (Theischinger and Hawking, 2006). pterostigma, Rspl originating more basally and a well-defined IR1 differs from Cretaeshna in having two rows of cells (Wilson and Xu, 2008). Any affinity between Cretaeshna and Pet- between RP3/4 and MA under Rspl instead of only one row in the aliaeschna can be excluded since the latter has many straight latter and a longer pterostigma (Theischinger and Hawking, 2006). intercalary veins between the main veins, a dense wing venation An affinity between Cretaeshna and Planaeschna can be excluded and a well-defined, long IR1 (Karube, 2000). since the latter has a well-defined IR1, a long pterostigma and a In conclusion, Cretaeshna is mostly like a member of Tele- basally originating Rspl (Wilson and Xu, 2008). phlebiinae but differs from the latter in having a weakly-defined 108 D. Zheng et al. / Cretaceous Research 72 (2017) 105e109

Fig. 3. Cretaeshna lini gen. et sp. nov., holotype, NIGP164824. Photographs showing details of pterostigmal area (A), nodal area (B), forewing apex (CeD).

IR1 and a short pterostigma. Other characters of Cretaeshna are 5. Conclusions shared with several members of Telephlebiinae. Here we propose a new genus for the unique type specimen, but more specimens with A new true dragonfly, Cretaeshna lini gen. et sp. nov., is described the basal part of the wing preserved would help to confirm and from mid-Cretaceous Burmese amber. This is the first aeshnid clarify this attribution. (hawker) dragonfly known as an amber inclusion. Our find D. Zheng et al. / Cretaceous Research 72 (2017) 105e109 109 augments the diversity of Mesozoic true dragonflies, and enhances McLachlan, R., 1896. On some Odonata of the subfamily Aeschnina. Annals and e our understanding of the palaeogeographic distribution of aeshnid Magazine of Natural History, series 6 17 (102), 409 425. Nel, A., Martinez-Delclos, X., Paicheler, J.C., Henrotay, M., 1993. Les ‘Anisozygoptera’ dragonflies, reported for the first time in the mid-Cretaceous of SE fossiles. 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