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From the Mid-Cretaceous Amber of Myanmar Cretaceous Research Cretaceous Research 78 (2017) 78e83 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes The first moth lacewing (Insecta: Neuroptera: Ithonidae) from the mid-Cretaceous amber of Myanmar * ** Xiumei Lu a, c, Weiwei Zhang b, Michael Ohl c, , Xingyue Liu a, a Department of Entomology, China Agricultural University, Beijing 100193, China b Three Gorges Entomological Museum, P.O. Box 4680, Chongqing 400015, China c Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitaetsforschung Invalidenstraße 43, Berlin 10115, Germany article info abstract Article history: The lacewing family Ithonidae is reported from the mid-Cretaceous amber of Myanmar for the first time. Received 12 February 2017 A new genus and species, Burmithone pennyi gen. et sp. nov., is herein described based on an almost Received in revised form completely preserved female specimen. The new genus exhibits a number of remarkable forewing 10 May 2017 characters, such as the proximal branches of RP vein fused with the MA vein and the peculiar configu- Accepted in revised form 25 May 2017 ration of MP and CuA. The systematic position of Burmithone gen. nov. is briefly discussed in comparison Available online 26 May 2017 with other genera of Ithonidae. © 2017 Elsevier Ltd. All rights reserved. Keywords: Neuropterida Ithonidae Taxonomy Burmese amber Mesozoic 1. Introduction veinlet, and the presence of nygmata. The larval stage of Ithonidae is unique because of its peculiar scarabaeiform (grub-like C-sha- Ithonidae sensu lato presents as an archaic small family of ped) older instar larvae (Grebennikov, 2004), which are subterra- Neuroptera, comprising the moth lacewings (Ithonidae sensu nean and suggested to be phytophagous (Gallard, 1932; Faulkner, stricto), montane lacewings (former Rapismatidae) and giant 1990). Both recent morphology-based and molecule-based phylo- lacewings (former Polystoechotidae) according to the recent genetic studies suggest a sister group relationship between Itho- taxonomic treatment, which placed the preceding three family- nidae and Myrmeleontiformia (Aspock€ and Aspock,€ 2008; group taxa into a single Ithonidae in the broad sense based on Winterton et al., 2010; Wang et al., 2017). total-evidence phylogenetic analysis (Winterton and Makarkin, Currently, Ithonidae include 10 extant genera and ca. 40 species 2010). Nonetheless, Zheng et al. (2016a) provided morphological worldwide. The earliest definite ithonid fossil is known from the evidence to divide Ithonidae sensu lato into three genus-groups, Middle Jurassic of China (Zheng et al., 2016a), and 11 genera and 28 i.e., the Ithonid genus-group, the Polystoechotid genus-group, fossil ithonids are recorded from the interval between the Middle and the Rapismatid genus-group, although the group members Jurassic to the late Eocene (Makarkin et al., 2014; Zheng et al., of the preceding three genus-groups are not equivalent to that 2016a,b). Among them, only one species was found in amber, i.e., placed in the former three families and include many fossil the late Eocene Baltic amber (Makarkin et al., 2014). genera. Here we report a new genus and species of Ithonidae from the Most adults of Ithonidae sensu lato are characterized by the mid-Cretaceous amber of Myanmar based on a nearly completed robust body, the head more or less retracted under the pronotum, female specimen. It represents the first record of Ithonidae from the broad forewing costal space with branched recurrent humeral this deposit, where a remarkably diverse paleofauna of Neuro- ptera was present (see Makarkin, 2016). Some noteworthy morphological characters in the new ithonid provide new insights * Corresponding author. for understanding the taxonomy and evolutionary history of ** Corresponding author. Ithonidae. E-mail addresses: [email protected] (M. Ohl), [email protected] (X. Liu). http://dx.doi.org/10.1016/j.cretres.2017.05.027 0195-6671/© 2017 Elsevier Ltd. All rights reserved. X. Lu et al. / Cretaceous Research 78 (2017) 78e83 79 2. Material and methods The amber sample under present study is from the Hukawng Valley in Tanai Township, Myitkyina District of Kachin State, Myanmar (see Kania et al., 2015: fig. 1). The age of this deposit has been investigated and dated to be ~99 MA (earliest Cenomanian) by UePb dating of zircons from the volcaniclastic matrix of the amber (Shi et al., 2012b). The type specimen is currently housed in the Entomological Museum, China Agricultural University (CAU), Beijing, and it will eventually be deposited in the Three Gorges Entomological Museum (EMTG), Chongqing (specimen available for study by contacting WZ). Photographs and drawings were taken and made using a Zeiss SteREO Discovery V12 stereo microscope system. The figures were prepared with Adobe Photoshop CS4. Terminology of wing venation generally follows H. Aspock€ et al. (1980) and Kukalova-Peck and Lawrence (2004). Terminology of genitalia fol- lows Aspock€ and Aspock€ (2008). Abbreviations used for wing veins are: A, anal vein; C, costa; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; M, media; MA, media anterior; MP, media posterior; R, radius; RA, radius anterior; RP, radius posterior; ScA, subcosta anterior; ScP, subcosta posterior. 3. Systematic palaeontology Order Neuroptera Linnaeus, 1758 Family Ithonidae Newman, 1853 sensu Winterton and Makarkin, 2010 Genus Burmithone gen. nov. Figs. 1e4 urn:lsid:zoobank.org:act:6F8A3661-B2AA-46C0-8C0D- 29D61C898C42 Type species: Burmithone pennyi sp. nov. Diagnosis. The new genus can be distinguished from the other species of Ithonidae by a combination of the following characters: (1) head completely retracted under a nearly hexagonal pronotum [partly protruding from pronotum in the Polystoechotid genus- group]; (2) antenna almost half of forewing length [very short in the Polystoechotid genus-group and most species of the Rap- ismatid and Ithonid genus-groups, being less than half of the forewing length]; (3) nygmata absent [present in most genera of Ithonidae]; (4) forewing recurrent humeral veinlet angulately curved [arcuately curved in most genera of Ithonidae except Prin- cipiala Makarkin & Menon and Allorapisma Makarkin & Archibald]; Fig. 1. Burmithone pennyi gen. et sp. nov., holotype female. A. Habitus photograph, (5) interlink veinlets among costal crossveins largely reduced in dorsal review; B. Habitus photograph, ventral review; C. Photograph of tarsi. Scale bar: forewing costal space [forewing costal space with several or many 2.5 mm (A, B); 1.0 mm (C). interlink veinlets among costal crossveins in many genera of the Rapismatid and Ithonid genus-groups]; (6) ScP stout, ending general irregularly arranged [regularly arranged into at least one sharply curved and fused with RA into ScP þ RA that reaches gradate series in the Polystoechotid genus-group]. anterior margin straightly before wing apex [ScP and RA distinctly Etymology. From Burma (¼ Myanmar) and Ithone (the type genus- separated in most genera of the Rapismatid and Ithonid genus- group name of Ithonidae), in reference to the occurrence of the groups]; (7) forewing RP with proximal two branches fused with new genus from the mid-Cretaceous amber of Myanmar. Gender: MA, forming two long cells [such fusion absent in most genera of Feminine. Ithonidae except Principiala and Allorapisma]; (8) forewing MP Remarks. Placement of the new genus in Ithonidae is undoubted lacking primary forking [deeply branched in most genera of Itho- based on the following features: robust body; head completely nidae except Principiala and Allorapisma ]; (9) forewing CuA pro- retracted under pronotum; proximal part of forewing costal space fusely branched anteriad from its midpoint, forming a very broad with branched recurrent humeral veinlet; presence of subtriangular area [less branched in most genera of Ithonidae numerous and irregularly arranged crossveins. except Principiala and Allorapisma, or densely branched but with branches directed posteriad in e.g. Megalithone Riek and Platys- The new genus could be placed in a suprageneric taxon, namely toechotes Carpenter (see Fig. 4)]; (10) forewing with crossveins in the Principiala group, proposed by Makarkin and Archibald (2009) 80 X. Lu et al. / Cretaceous Research 78 (2017) 78e83 Fig. 2. Burmithone pennyi gen. et sp. nov., holotype female. A. Drawing of right forewing; B. Drawing of left forewing. Scale bar: 1.0 mm. by the proximal branches of RP fused with MA and the peculiar Diagnosis. As for the genus. configuration of MP and CuA, which is however interpreted Description. Female. Body length 7.91 mm; head 0.80 mm long and differently in this paper (see Discussion). The Principiala group 1.93 mm wide; antenna length 5.36 mm; pronotum 1.50 mm long previously comprised two fossil genera, i.e. Principiala from the and 2.40 mm wide; forewing 8.89 mm long and 4.20 mm wide; Lower Cretaceous of Brazil and England and Allorapisma from the abdomen length 4.42 mm. Foreleg tarsus length: 0.81 mm; claw early Eocene of U.S.A. Obviously, the new genus, Burmithone gen. length 0.10 mm; midleg claw length 0.15 mm; hind leg claw length nov. is closely related to Principiala and Allorapisma based on the 0.08 mm. unique feature of forewing venation within Ithonidae mentioned above. However, Burmithone gen. nov. can be distinguished from Head wide, but slightly thinner than pronotum; compound eyes Principiala by the relatively long antenna (5.36 mm long) [rather globular; antenna thickly filiform, nearly half of forewing
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