See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/297518833 †Alienoptera - A new insect order in the roach- mantodean twilight zone Article in Gondwana Research · March 2016 DOI: 10.1016/j.gr.2016.02.002 CITATIONS READS 12 1,036 6 authors, including: Ming Bai Rolf Beutel Chinese Academy of Sciences Friedrich Schiller University Jena 127 PUBLICATIONS 374 CITATIONS 287 PUBLICATIONS 6,272 CITATIONS SEE PROFILE SEE PROFILE Klaus-Dieter Klass Benjamin Wipfler Senckenberg Research Institute Friedrich Schiller University Jena 76 PUBLICATIONS 1,451 CITATIONS 60 PUBLICATIONS 931 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Insect Systematics and Evolutionary Biology Forum View project Genitalic region in Archaeognatha, Zygentoma and Odonata: comparative morphology, functional reconstructions, phylogenetic value, and evolution View project All content following this page was uploaded by Rolf Beutel on 04 April 2016. The user has requested enhancement of the downloaded file. GR-01582; No of Pages 10 Gondwana Research xxx (2016) xxx–xxx Contents lists available at ScienceDirect Gondwana Research journal homepage: www.elsevier.com/locate/gr †Alienoptera — A new insect order in the roach–mantodean twilight zone Ming Bai a, Rolf Georg Beutel b,⁎, Klaus-Dieter Klass c,WeiweiZhanga,d,XingkeYanga,BenjaminWipfler a,b a Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District, Beijing 100101, China b Entomology Group, Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, Friedrich-Schiller-Universität Jena, Erbertstrasse 1, D-07743 Jena, Germany c Senckenberg Natural History Collections Dresden, Königsbrücker Landstrasse 159, D-01109 Dresden, Germany d P.O. Box 4680, Chongqing 400015, China article info abstract Article history: A new insect species (†Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang gen. et sp. nov.) of a new Received 20 November 2015 order and family is described, based on a single male embedded in Cretaceous Burmese amber (ca. 99 Ma). Un- Received in revised form 29 January 2016 usual characters are shortened forewings combined with fully developed, operational hindwings, similar as in Accepted 2 February 2016 Dermaptera, and specialized attachment pads otherwise only found in mantophasmatodeans (heelwalkers). A Available online xxxx cladistic analysis suggests a placement as sister to Mantodea, supported by a profemoral brush and other charac- fl Handling Editor: I.D. Somerville ters. The male genitalia show the same pattern in both groups. Specialized features are the unusual ight appa- ratus, attachment structures adapted for locomotion on leaves, and a dense profemoral setation suitable for Keywords: catching small prey. †Alienopterus was apparently able to fly and likely a predator of small arthropods in bushes Alienoptera or trees. An impressive radiation of Mantodea started in similar habitats at least 35 Ma later in the early Cenozoic. New order In contrast, †Alienopterus was an evolutionary dead end in the roach–mantis transition zone. Insecta © 2016 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved. Mantodea Amber Cretaceous 1. Introduction distinctly shortened ovipositor distinguishes “modern” dictyopterans from the “roachoids”, which persisted up to the end of the Mesozoic In this study we describe a new extinct insect species in a new (Grimaldi and Engel, 2005; Legendre et al., 2015). monotypic order. Its potential role as a link between roach-like insects In the typical case roaches are saprophagous and solitary with a and the praying mantises is evaluated. The single specimen with a high- ground-oriented lifestyle (Bell et al., 2007), and the same is likely true ly unusual character combination is embedded in Cretaceous Burmese for most of the “roachoids”. However, during their history of more amber, which has turned out as a very rich source of insect fossils than 300 Ma, more or less highly specialized groups have evolved sever- (Grimaldi et al., 2002). al times, for example the Mesozoic †Umenocoleidae, a group strongly The earliest roach-like insects known from the Upper Carboniferous resembling beetles, with heavily sclerotized tegmina and a small were already remarkably diverse (Labandeira, 1994; Grimaldi and pronotum (Vršanský, 2003; Grimaldi and Engel, 2005; Nel et al., Engel, 2005; Legendre et al., 2015). Similar to extant cockroaches, 2014), or the Jurassic and Cretaceous †Raphidiomimidae (Liang et al., Palaeozoic forms with a preserved body had a discoid pronotum partly 2009), predacious forms with raptorial forelegs (Grimaldi and Ross, covering the head, flattened posteriorly slanting coxae, and leathery 2004). forewings (tegmina). In Carboniferous coal swamps they were abun- In contrast to these lineages, which vanished from the scene in the dant and arguably the dominant group of insects (Grimaldi and Engel, late Mesozoic, “modern” Dictyoptera are a quite successful group with 2005). The phylogenetic relationships among these Palaeozoic almost 10,000 described species. Aside from the more or less general- “roachoids” are unresolved as yet (Legendre et al., 2015). However, it ized roaches (“Blattaria”), the group contains two highly specialized lin- is widely accepted that one of these lineages gave rise to Dictyoptera, eages: the wood- or fungus-feeding and eusocial termites (Isoptera), most likely in the Jurassic (Grimaldi and Engel, 2005: fig. 7.60; Misof today recognized as a subordinate group of Blattodea (Klass and et al., 2014; Legendre et al., 2015; Tong et al., 2015). This clade com- Meier, 2006; Inward et al., 2007), and the predacious praying mantises prises the “modern” roaches (Blattodea) including termites (Isoptera) (Mantodea), which are characterized by specifically modified raptorial and the praying mantises (Mantodea) (Grimaldi, 2003; Grimaldi and forelegs and an elaborate prey catching mechanism. With about 3000 Engel, 2005; Legendre et al., 2015). Despite a general similarity, the and 2300 extant species (Ehrmann, 2002; Krishna et al., 2013), respec- tively, termites and praying mantises are slightly less species-rich than “ ” ⁎ Corresponding author. their generalized relatives, the paraphyletic roaches ( Blattaria ) E-mail address: [email protected] (R.G. Beutel). (Beutel et al., 2014). http://dx.doi.org/10.1016/j.gr.2016.02.002 1342-937X/© 2016 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved. Please cite this article as: Bai, M., et al., †Alienoptera — A new insect order in the roach–mantodean twilight zone, Gondwana Research (2016), http://dx.doi.org/10.1016/j.gr.2016.02.002 2 M. Bai et al. / Gondwana Research xxx (2016) xxx–xxx Looking at extant species, Mantodea can be easily recognized by and Odonata were added as outgroup terminals(Kristensen, 1991; conspicuous modifications linked with the specialized predacious life Beutel and Gorb, 2006; Misof et al., 2014), without enforcing the mono- style. Even the few known early-diverging extinct forms (Grimaldi, phyly of the ingroup. Additionally, we included three extinct roaches 2003) differ distinctly from non-mantodean dictyopterans, including which have been considered as raptorial (†Ponopterix, †Jantaropterix those considered as predators (Grimaldi and Ross, 2004; Hörnig et al., and †Manipulator). As mantodean terminals we included four extant 2013; Lee, 2014; Vršanský and Bechly, 2015). In the present study we genera (Chaeteessa, Metallyticus, Hymenopus and Stagmomantis)and describe and document a Cretaceous fossil in detail using modern tech- all described fossils with at least some accessible characters niques. The morphological features are evaluated with respect to the (†Burmantis, †Jersimantis, †Santanmantis and †Ambermantis). Supple- phylogenetic position. Implications for the life style and habitats of the mentary Method S1 provides detailed information about the studied extinct species are discussed. An evolutionary scenario is developed taxa including the source of information. The 58 characters (from based on the phylogenetic conclusions, with a special focus on character head, mouthparts, pronotum, pterothorax, wings, legs, and transformations in the “roach–mantis transition zone”. postabdomen) are listed in Supplementary Method S2. To address the relationships of major lineages a few selected characters were included 2. Material and methods which cannot be scored for fossil terminals (e.g., internal parts of the exoskeleton, muscles, mode of insemination). We analyzed one matrix 2.1. Material and photography (Supplementary Method S3) with all taxa included and a second one (Supplementary Method S4) with fossil taxa with many inaccessible The single specimen, a male, was obtained from an amber deposit in characters excluded (fossil roaches and mantodean nymphs). The data the Hukawng Valley of Myanmar. The age has been estimated as were gathered in a matrix with WinClada and parsimony analyses ca. 99 Ma (98.8 ± 0.6; earliest Cenomanian) based on U–Pb dating of were conducted with NONA (ratchet, 1000 replicates) and TNT (tradi- zircons from the volcaniclastic matrix of the amber (Shi et al., 2012). tional search; 99,999 random seeds, 1000 replicates) (Goloboff et al., The mining locality is at Noije Bum, near Tanai Village (26°21′33.41″N, 2003, 2008). Bremer