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New Stem-Phasmatodea from the Middle Jurassic of China

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Eur. J. Entomol. 108: 677–685, 2011 http://www.eje.cz/scripts/viewabstract.php?abstract=1667 ISSN 1210-5759 (print), 1802-8829 (online) New stem-Phasmatodea from the Middle Jurassic of China LIANGJIE SHANG1, OLIVIER BÉTHOUX2 and DONG REN1* 1College of Life Science, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China; e-mail: [email protected] 240 rue d’Aveillans, 38770 La Motte d’Aveillans, France; e-mail: [email protected] Key words. Adjacivena rasnitsyni gen. n., sp. n., Phasmatodea, Susumaniidae, Heteropteryx dilatata, China, Daohugou, intra-specific variability, sexual dimorphism, fossil, Middle Jurassic Abstract. Adjacivena rasnitsyni gen. n., sp. n., from the Middle Jurassic Jiulongshan Formation (Daohugou Village, Inner Mongolia, China), is described based on a set of well-preserved specimens, interpreted as two females and a male of the same species. The new taxon is assigned to the family Susumaniidae, i.e. is a candidate stem-Phasmatodea. This ordinal assignment is supported by genital elements, chiefly a putative operculum concealing the ovipositor observed in one female specimen and a putative extension of the 10th tergum with a thorn pad in the male specimen. Variation in the venation of the forewing is appreciated based on more or less complete pairs of wings and a comparison with that in the forewings of male Heteropteryx dilatata (Parkinson, 1798). This material offers new perspectives on the primary homologies of the hind wing venation of stick-insects. INTRODUCTION “Cu” is concave). This situation has two interrelated con- Stick-insects (also known as Cheleutoptera, Phasmato- sequences: conjectures about the primary homologies of dea, Phasmida and Phasmatoptera, among others) com- stick-insect wing venation and the identification of sev- pose a rather small order of insects, which currently eral Mesozoic representatives of the order are both uncer- includes over 3,000 extant species (Bragg, 2001; Brock & tain. Hasenpusch, 2009). The comparatively low extant diver- Thus, specimens of Mesozoic fossils on which the sity in this order is mirrored by its rarity in the fossil details of the hind wings and body are clearly visible are record: a few tens of species are to varying extents con- particularly valuable. Herein we describe three new and vincingly assigned to this order (Birket-Smith, 1981; comparatively complete Jurassic specimens from China Gorochov, 1988, 1992, 1993, 1994, 2000; Martynov, that we tentatively assign to Phasmatodea. 1928; Nel et al., 2004, 2010; Ren, 1997; Sharov, 1968, MATERIAL AND METHODS 1971; Tillyard, 1918; Wedmann et al., 2007; among All the fossil material described is housed in the Key Labora- others). However actual affinities of many fossil taxa tory of Insect Evolution & Environmental Changes, Capital remain uncertain (Grimaldi & Engel, 2005; Tilgner, 2000; Normal University, Beijing, China. The material was collected and see discussion by Gorochov, 2000; Bradler & Buck- near Daohugou Village, Jiulongshan Formation, Ningcheng ley, 2011, with respect to Nel et al., 2010; and Delclòs et County, Inner Mongolia, China. The age of the Formation is al., 2008). still controversial, but most published biostratigraphic correla- This situation is primarily due to the incompleteness of tions and radiometric dates indicate a Middle Jurassic age the fossil material, but also to a lack of a clear-cut (Bathonian – Callovian boundary; Ren et al., 1995, 2002, 2010; forewing venation apomorphy for this order. According Cui et al., 2010; Zhao et al., 2010). to Gorochov & Rasnitsyn (2002), “fore wing with veins Fossil specimens were examined under a Leica MZ12.5 dis- secting microscope and illustrations made with the aid of a subparallel” is the only available character for identifying drawing tube attached to the microscope. Line drawings were a stick-insect wing. However, this character is shared by a prepared using CorelDraw 12 graphic software. Photographs large number of narrow-winged polyneopteran insects, were taken using a Nikon D100 coupled to a 105 mm Nikon including, for example, some mantids. In addition the fact macro lens and processed using Adobe Photoshop. that many stick-insects have vestigial forewings reduces Material of Heteropteryx dilatata (Parkinson, 1798) was the value of this character. Finally, although relevant at obtained from an insect dealer. According to the data provided the level of the order, the comparative analysis of fore- the specimens were collected from wild populations at Tapah, and hind wing venation provided by Ragge (1955), based Cameron Highlands (Malaysia) in February 2006. Wings were on extant material, needs to be reinterpreted in a broader cut off and mounted on slides in Euparal medium. Photographs context, chiefly because the CuA vein in the hind wings, were taken using a Canon EOS 400D coupled to a 50 mm Canon macro lens and processed using Adobe Photoshop. which is strongly convex in nearly all polyneopteran The systematic arrangement tentatively follows Gorochov insects (possibly nearly all winged insects), is not (1988, 2000; but see below). Except “Sc” and “RS” are replaced included in this interpretation (the vein interpreted as * Corresponding author. 677 by “ScP” and “RP”, respectively, wing venation nomenclature Gorochov (2000) is the location of the first fork of RP in and conjectures of primary homology follow Sharov (1968, forewings, which he states is located “in proximal wing 1971; followed by Carpenter, 1992). This does not imply sup- part” in Susumaniinae and “in middle wing part” in Phas- port for this primary homology by the second author. Wing momimoidinae. However, according to data published by venation abbreviations are as follows: ScP – posterior Subcosta; Birket-Smith (1981: Fig. 1D) and Gorochov (2000: Fig. R – Radius; RA – anterior Radius; RP – posterior Radius; MA – anterior Media; MA1 – anterior branch of MA; MA2 – posterior 3c) this fork is located about 40.6% along the length of branch of MA; MP – posterior Media; CuA – anterior Cubitus; the wing in Coniphasma rosenkrantzi Birket-Smith, 1981 CuA1 – anterior branch of CuA; CuA2 – posterior branch of [a Susumaniinae according to Gorochov (2000)] and CuA; CuP – posterior Cubitus; 1A – first anal vein. Other abbre- 41.5% along the length of the wing in Phasmomimoides viations and indications are as follows: LFW – left forewing; lineatus Sharov, 1968 (type-species of the type-genus of RFW – right forewing; LHW – left hind wing; RHW – right the Phasmomimoidinae). The difference is therefore neg- hind wing; on Fig. 3, * indicate the claval fold on each ligible. Because of this and our limited knowledge of the forewing. Abbreviations for genital structure follow Bradler intra-specific variation in wing venation in Susumaniidae, (2009). In detail: Gon – gonobasis; Op – operculum; Gap8 – we propose to compare the material of Adjacivena rasnit- gonapophysis 8; Cer – cercus. syni gen. n., sp. n. with all taxa assigned to the Susu- SYSTEMATICS maniidae by Gorochov (2000). Palaeopteron complexus Rice, 1969 is documented Order Phasmatodea Brunner von Wattenwyl, 1893 based on a single specimen, several wings of which over- Superfamily Susumanioidea Gorochov, 1988 lap. Only a part of the hind wing venation is deciphered Family Susumaniidae Gorochov, 1988 by Rice (1969). According to the available data the stem Genus Adjacivena gen. n. of RP + MA1 [“R2+3” & “R4+5” according to Rice Type species. Adjacivena rasnitsyni sp. n. by present descrip- (1969)] is comparatively long and MA2 (“M1+2”) and tion (monotypic genus). MP + CuA1 (“M3+4”) reach CuA2, which is similar to what occurs in Orephasma eumorpha Ren, 1997, and Composition. Type species and Adjacivenia reticulata make Adjacivena rasnitsyni gen. n., sp. n. distinct from (Kuzmina, 1985) comb. n. (but see discussion; originally these species. assigned to the genus Phasmomimoides Sharov, 1968, In Adjacivena rasnitsyni gen. n., sp. n. MA2 and MP + and then to the genus Eosusumania Gorochov, 1988 by CuA1 are very narrowly separated over some distance. Gorochov, 1988). This trait is very probably derived and is absent in Coni- Diagnosis. Forewing: MA2 approaching MP + CuA1 a phasma rosenkrantzi (Birket-Smith, 1981). short distance before the middle of the wing (main diag- The interpretation of the forewing venation of Promas- nostic character); MP + CuA1 forked (known only for tacoides albertae Kevan & Wighton, 1981 is not evident type species). Hind wing (known only for type species): from the available data (Kevan & Wighton, 1981: Fig. 6): MA1 fused over a moderate distance with RP; MA2 fused the posterior branch of the vein interpreted by Kevan & with MA1 distal to its divergence from RP + MA1. Legs Wighton (1981) as “MA” together with “MP” probably slender, moderately long and apparently not armed. form MP + CuA1; the anterior branch of the vein inter- Etymology. Generic name derived from the combination of preted as “MA” is probably MA2; and the posterior “adjac-” (adjacent in Latin) and “vena” (vein in Latin) referring branch of that interpreted as “Rs”, the origin of which is to the main character of this genus, namely that the vein MA2 unclear, is probably “MA1”. It is plausible that there are approaches MP + CuA1 in the forewings. Gender: feminine. secondary intercalary veins in the area between the two Discussion. The occurrence of a RP vein in the branches of RP and in that between the posterior branch forewing that forks near its origin is an uncommon trait in of RP and MA1. These are absent in our material, in winged insects. This trait is present in the forewing of which the cross-veins are reticulated. In addition the col- male Heteropteryx dilatata (extant Phasmatodea; Will- oration pattern differs significantly. mann, 2003: Fig. 12; Fig. 4) and in several genera Phasmomimella paskapoensis Kevan & Wighton, 1981 assigned to the family Susumaniidae (see composition in is only known from a hind wing. In the light of recent dis- Gorochov, 2000). Affinities of the new material with coveries (Ren, 1997; and herein) the posterior branch of Blattinopsidae Bolton, 1925, which also have a RP with “Rs”, as indicated by Kevan & Wighton (1981: Fig.
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    BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 20 THE EARWIGS OF CALIFORNIA (Order Dermaptera) BY ROBERT L. LANGSTON and J. A. POWELL UNIVERSITY OF CALIFORNIA PRESS THE EARWIGS OF CALIFORNIA (Order Dermaptera) BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 20 THE EARWIGS OF CALIFORNIA (Order Dermaptera) BY ROBERT L. LANGSTON and J. A. POWELL UNIVERSITY OF CALIFORNIA PRESS BERKELEY LOS ANGELES LONDON 1975 BULLETIN OF THE CALIFORNIA INSECT SURVEY Advisory Editors: H. V. Daly, J. A. Powell; J. N. Belkin, R. M. Bohart, R. L. Doutt, D. P. Furman, J. D. Pinto, E. I. Schlinger, R. W. Thorp VOLUME 20 Approved for publication September 20,1974 Issued August 15, 1975 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND ISBN 0-520-09524-3 LIBRARY OF CONGRESS CATALOG CARD NUMBER: 74-22940 0 1975 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA CONTENTS Introduction .................................................. 1 California fauna ............................................. 1 Biology ................................................... 1 History of establishment and spread of introduced species in California ........ 2 Analysis of data ............................................. 4 Acknowledgments ............................................ 4 Systematic Treatment Classification ............................................... 6 Key to California species ........................................ 6 Anisolabis maritima (Ght5) ...................................
  • Insect Egg Size and Shape Evolve with Ecology but Not Developmental Rate Samuel H

    Insect Egg Size and Shape Evolve with Ecology but Not Developmental Rate Samuel H

    ARTICLE https://doi.org/10.1038/s41586-019-1302-4 Insect egg size and shape evolve with ecology but not developmental rate Samuel H. Church1,4*, Seth Donoughe1,3,4, Bruno A. S. de Medeiros1 & Cassandra G. Extavour1,2* Over the course of evolution, organism size has diversified markedly. Changes in size are thought to have occurred because of developmental, morphological and/or ecological pressures. To perform phylogenetic tests of the potential effects of these pressures, here we generated a dataset of more than ten thousand descriptions of insect eggs, and combined these with genetic and life-history datasets. We show that, across eight orders of magnitude of variation in egg volume, the relationship between size and shape itself evolves, such that previously predicted global patterns of scaling do not adequately explain the diversity in egg shapes. We show that egg size is not correlated with developmental rate and that, for many insects, egg size is not correlated with adult body size. Instead, we find that the evolution of parasitoidism and aquatic oviposition help to explain the diversification in the size and shape of insect eggs. Our study suggests that where eggs are laid, rather than universal allometric constants, underlies the evolution of insect egg size and shape. Size is a fundamental factor in many biological processes. The size of an 526 families and every currently described extant hexapod order24 organism may affect interactions both with other organisms and with (Fig. 1a and Supplementary Fig. 1). We combined this dataset with the environment1,2, it scales with features of morphology and physi- backbone hexapod phylogenies25,26 that we enriched to include taxa ology3, and larger animals often have higher fitness4.