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Stick Insect in Burmese Amber Reveals.Pdf Downloaded from http://rspb.royalsocietypublishing.org/ on April 25, 2018 Stick insect in Burmese amber reveals rspb.royalsocietypublishing.org an early evolution of lateral lamellae in the Mesozoic Sha Chen1, Xiangchu Yin2, Xiaodan Lin1, Chungkun Shih1,3, Runzhi Zhang4, Research Taiping Gao1 and Dong Ren1 Cite this article: Chen S, Yin X, Lin X, Shih C, 1College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, Zhang R, Gao T, Ren D. 2018 Stick insect in People’s Republic of China Burmese amber reveals an early evolution of 2Northwest Institute of Plateau Biology, Chinese Academy of Sciences, 23, Xinning Road, 810008 Xining, People’s Republic of China lateral lamellae in the Mesozoic. Proc. R. Soc. B 3Department of Paleobiology, Smithsonian Institution, National Museum of Natural History, Washington, 285: 20180425. DC 20013-7012, USA http://dx.doi.org/10.1098/rspb.2018.0425 4State Key Laboratory of Integrated Management of Pest Insects and Rodents, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, People’s Republic of China DR, 0000-0001-8660-0901 Received: 24 February 2018 Extant stick and leaf insects commonly imitate twigs or leaves, with lateral Accepted: 28 March 2018 lamellae used to enhance crypsis or achieve mimicry for protection. However, the origin and early evolution of such lateral expansions among Phasmatodea are unknown, because all known Mesozoic phasmatodeans hitherto lack preserved evidence of such structures. We report here the first Mesozoic stick insect, Elasmophasma stictum gen. et sp. nov., with well-preserved, thin, Subject Category: lateral lamellae on the thoracic pleura, the terga of abdominal segments I–X Palaeobiology and the ventrolateral margins of all femora. This new species, from the mid- Cretaceous amber of northern Myanmar, has a clear, stick-like body and is Subject Areas: assigned to Euphasmatodea. The abdominal structures of E. stictum exhibit behaviour, evolution, palaeontology traces of multiple expansions of the terga, suggesting that such structure might have been an early development of body expansions used to improve Keywords: crypsis for stick or leaf insects when they sprawled on twigs or leaves. Elasmophasma, Euphasmatodea, lamella, mimicry, crypsis, Myanmar 1. Introduction Phasmatodea, stick and leaf insects, are a rather small insect order, comprising Authors for correspondence: approximately 3000 described extant species [1,2]. Phasmatodeans typically live Taiping Gao on trees or shrubs throughout the world, especially in the tropics and subtro- e-mail: [email protected] pics [3–5]. Most extant stick insects imitate the shape and colour of thin Dong Ren twigs, but some stick insects in the genera Cotylosoma and Extatosoma (Phasma- e-mail: [email protected] tidae) (figure 1a,b) have abdominal lobes expanded from terga and/or legs as mimicry of leaves to improve crypsis. Living leaf insects (Phylliidae), compris- ing only 1% of modern phasmatodean diversity, imitate leaves by means of specialized expansive structures on the legs, thorax, wings and abdomen [6]. Many fossil ‘stick insects’ have been described from the Mesozoic, and most of these species were erected based on isolated wings or wing fragments. Owing to significant differences between Mesozoic and extant stick insects, the classification and relationships among these groups remain contentious [7,8]. Furthermore, none of the Mesozoic lineages show evidence for the lateral body expansions associated with adaptive crypsis, instead exhibiting elongate forms [8,9]. Two mid-Cretaceous species, Pseudoperla scapiforma and P. leptoclada from Myanmar, have elongate bodies and slightly curved profe- mora, but aside from their superficial form relative to sticks, nothing more Electronic supplementary material is available could be concluded regarding potential mimicry [10]. online at https://dx.doi.org/10.6084/m9. Here we describe from the mid-Cretaceous amber of northern Myanmar a well-preserved, slender stick insect with thin, lateral lobe-like structures figshare.c.4061768. expanded from the body and femora. & 2018 The Author(s) Published by the Royal Society. All rights reserved. Downloaded from http://rspb.royalsocietypublishing.org/ on April 25, 2018 (a) (c) 2 rspb.royalsocietypublishing.org Proc. R. Soc. B (b) (d) 285 : 20180425 Figure 1. (a,b) Photographs of Extatosoma tiaratum Macleay, 1826 by Runzhi Zhang at Durban, South Africa on 10 July 2008. (c,d) E. stictum gen. et sp. nov. Holotype CNU-PHA-MA2017004. (c) Photograph of lateral view as preserved. (d) Line drawing of lateral view. Scale bars: (a,b) 10 mm and (c,d) 2 mm. 2. Material and methods pronotum. Metanotum and abdominal tergum I separated. Legs long, nearly equal to the length of body. Metafemora The amber type specimen described in this paper is housed in just similar to the other femora. Area apicalis present on all the Key Lab of Insect Evolution and Environmental Changes, tibiae and without spines. Femora, thorax and abdomen College of Life Sciences, Capital Normal University, Beijing, China (CNU; Dong Ren, Curator). with lateral expanded lamellae. Abdominal tergum X undiv- The specimen CNU-PHA-MA2017004 is preserved in a round ided. Cerci straight, undivided, circular in cross section, triangular piece of yellow amber, about 40 mm in maximum without prominence at base or thorns. length, 35 mm in maximum width and 10 mm in maximum thick- Remarks. The current classification of Phasmatodea is ness. The amber specimen was studied under a Leica M205C confusing [7,11–13]. Two broad groups, ‘Areolatae’ and ‘Anar- dissecting microscope. The habitus photographs were taken with a eolatae’, distinguished from each other based on the presence or Nikon SMZ 25 microscope with a Nikon DS-Ri 2 digital camera absence of the area apicalis, were historically recognized as two system, while the magnified images of details of the specimen were main clades, which are now known to be polyphyletic taken using a Nikon ECLIPSE Ni microscope with a Nikon DS-Ri 2 [11,12,14–16]. Elasmophasma stictum gen. et sp. nov. shares apo- digital camera system. Line drawing was prepared by using Adobe morphies with Phasmatodea as labrum emarginate [11,17,18], ILLUSTRATOR CC and Adobe PHOTOSHOP CC graphics software. prothorax with defensive glands [5,11,18,19], metasternum and abdominal sternum I fused [11,20], and cerci unsegmented [6,11,16,19,21]. Elasmophasma stictum sp. nov. can be further 3. Results assigned within Euphasmatodea based on the following Systematic palaeontology apomorphies: profurca largely reduced [5,11,17,18,21,22], pro- Order Phasmatodea Jacobson & Bianchi, 1902 and mesospina absent [5,11,23], profemur distinctly curved Suborder Euphasmatodea Bradler, 1999 basally [11,18] and tarsus with five articles [5,11,21,23], Family Incertae sedis although five-segmented tarsi are generally considered to be Elasmophasma gen. nov. plesiomorphic among Polyneoptera [1]. In addition, the Type species. Elasmophasma stictum sp. nov. antenna of E. stictum is much longer than body and foreleg, Etymology. The new generic name is a combination of the which is uncommon in stick insects. However, because the Greek word of elasmos, meaning ‘plate’, and phasma, meaning amber specimen is immature and lacks some diagnostic details, ‘ghost’, stem of the ordinal name of Phasmatodea. The name it cannot be placed further within that group and is deemed as refers to the lateral lamellae of the legs, thorax and abdomen. family Incertae sedis. This systematic relationship is consistent The gender of the name is neuter. with our phylogenetic analysis documented and presented in Diagnosis. Antenna much longer than body and foreleg. the electronic supplementary material. Cervix with a median plate. Prosternum with paired sensory The new species has seven annuli in the flagellum, and each areas. Mesonotum more than three times as long as flagellomere is greatly elongate. The eighth abdominal sternum Downloaded from http://rspb.royalsocietypublishing.org/ on April 25, 2018 (a) (c) 3 T8 rspb.royalsocietypublishing.org T9 T10 Ce Proc. R. Soc. B (b) Ta4 Ta2 Ta1 Un Ep Ar Ta3 Ta5 285 (d)(e)(f ) Pmx (g) : 20180425 Lb Md Af Prt Pgl Sc Mst Pd Plb Gl Fla I Mst Cerv Mtt Apg T1 Mtt Tr Cx Bsl Figure 2. Elasmophasma stictum gen. et sp. nov. Photographs of Holotype CNU-PHA-MA2017004. (a) Lateral view as preserved. (b) Left mesotarsus in lateral view. (c) T8–10, epiproct and cerci in dorsal view. (d) Mesonotum, metanotum and the first abdominal tergum. (e) Thoracic sterna. (f) Head in lateral view. (g) The area apicalis (arrows) of right protibia in the ventral view. Scale bars: (a) 2 mm; (b–f) 0.2 mm and (g) 0.1 mm. Af, antennal field; Apg, aperture of the pronotal gland; Ar, arolium; Bs1, prothoracic basisternum; Ce, cercus; Cerv, cervix; Cx, coxa; Ep, epiproct; Fla I, flagellomere I; Gl, glossa; Lb, labrum; Md, mandible; Mst, mesothorax; Mtt, metathorax; Pd, pedicellus; Pgl, paraglossa; Plb, labial palpus; Pmx, maxillary palpus; Prt, prothorax; Sc, scape; T1, the first abdominal tergum (median segment); T8–T10, abdominal tergum 8–10; Ta1–Ta5, tarsomere I–V; Tr, trochanter; Un, claw. lacks paired tubercles and a transverse fold medially, but Holotype. A male first-instar nymph, no. CNU-PHA- the ninth abdominal sternum has a slight mounding, which MA2017004, deposited in Capital Normal University, Beijing. suggest that the new specimen may be a male first-instar Type locality and horizon. The amber specimen was collected nymph [24]. The new genus is differentiated from other from Kachin (Hukawng Valley) of northern Myanmar, which groups of Euphasmatodea mainly by the three characters was dated at 98.79 + 0.62 Ma [26,27]. femora, mesothorax and metathorax, and abdominal segments Description. Bodyelongate, slender (figures 1c,d and 2a), with I–X with lateral expanded lamellae; prosternum with paired a longitudinal median carina in ventral view; surface smooth. sensory areas and metanotum separated from the first abdomi- Head subglobose, longer than wide; ocelli absent; compound nal tergum.
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