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Fossil Record of Sternorrhyncha (Hemiptera)

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Fossil Record of Sternorrhyncha (Hemiptera) Bo Wang 1, Jacek Szwedo 2 1 Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, China; e-mail; [email protected] 2 Department of Invertebrate Zoology and Parasitology, University of Gdańsk, Poland; e-mail: [email protected] 2016 XXV International Congress of Entomology Orlando, Florida, USA | September 25-30 535: Symposium: Synthesis in Sternorrhyncha Systematics HYPOPERLIDA other Psocoptera Psocodea Liposcelidae Phthiraptera AMBLYCERA ISCHNOCERA RHYNCOPTHIRINA ANOPLURA HOLOMETABOLA PERMOPSOCIDA THRIPIDA PALEORRHYNCHA STERNORRHYNCHA Hemiptera FULGOROMORPHA CICADOMORPHA COLEORRHYNCHA HETEROPTERA Where to search for sternorrhynchan ancestors? What constitutes a sternorrhynchan insect? The recent morphological disparity and biodiversity is enormous, then which characters can be used for the fossils? Which is the oldest Sternorrhyncha? The questions where and when the splitting of Sternorrhyncha from hemipteran stock took place are not fully answered. INTERNATIONAL CHRONOSTRATIGRAPHIC CHART www.stratigraphy.org International Commission on Stratigraphy v 2016/04 numerical numerical numerical Eonothem numerical Series / Epoch Stage / Age Series / Epoch Stage / Age Series / Epoch Stage / Age Erathem / Era System / Period GSSP GSSP age (Ma) GSSP GSSP Eonothem /Eon Erathem /Era System /Period Eonothem /Eon Erathem /Era System /Period age (Ma) Eonothem /Eon Erathem /Era System /Period age (Ma) / Eon GSSA age (Ma) present ~ 145.0 358.9 ± 0.4 541.0 ±1.0 y Holocene r Ediacaran 0.0117 Tithonian a Upper 152.1 ±0.9 Famennian ~ 635 n Neo- Cryogenian r 0.126 Upper Kimmeridgian Middle Upper proterozoic ~ 720 e 157.3 ±1.0 t Pleistocene 0.781 372.2 ±1.6 a Calabrian Oxfordian Tonian u 1.80 163.5 ±1.0 Frasnian Callovian 1000 n c 166.1 ±1.2 Q Gelasian 2.58 i 382.7 ±1.6 Stenian Bathonian a s Piacenzian Middle 168.3 ±1.3 i s Bajocian Givetian 1200 Pliocene 3.600 170.3 ±1.4 n Meso- a 387.7 ±0.8 o Middle Ectasian Zanclean r Aalenian c v proterozoic 5.333 i u 174.1 ±1.0 Eifelian 1400 e o Messinian J 393.3 ±1.2 e Toarcian z 7.246 D Calymmian n 182.7 ±0.7 o r 1600 e Tortonian Emsian 11.63 c i e g Pliensbachian t Statherian o Lower o Serravallian 407.6 ±2.6 190.8 ±1.0 Lower o 13.82 z 1800 r e Miocene Pragian 410.8 ±2.8 n o c i Langhian Sinemurian P N a Orosirian 15.97 s i o 199.3 ±0.3 Lochkovian r Paleo- e z Burdigalian Hettangian 2050 201.3 ±0.2 419.2 ±3.2 b proterozoic o 20.44 M Rhyacian m n Aquitanian Rhaetian Pridoli a e 23.03 ~ 208.5 423.0 ±2.3 2300 Ludfordian c C 425.6 ±0.9 Chattian Ludlow e Siderian n 28.1 Gorstian 427.4 ±0.5 r Oligocene Upper Norian a 2500 P i c Homerian r Rupelian i Wenlock 430.5 ±0.7 Neo- u s ~ 227 33.9 l Sheinwoodian 433.4 ±0.8 archean i s a 2800 Priabonian Carnian S e i Telychian 37.8 r ~ 237 n n Llandovery 438.5 ±1.1 Meso- Bartonian T a e 41.2 Ladinian Aeronian 440.8 ±1.2 archean c c c e g ~ 242 i Eocene i Middle i Rhuddanian c h o Lutetian 3200 o o o 443.8 ±1.5 Anisian i c e z z z 247.2 o Hirnantian l 47.8 445.2 ±1.4 r Paleo- o o o z a Olenekian 251.2 A r r Lower r Ypresian o Katian archean P e e Induan 252.17 ±0.06 e Upper 56.0 e l n n n 3600 Changhsingian 254.14 ±0.07 453.0 ±0.7 n a a Thanetian a a 59.2 Lopingian Sandbian Eo- a h h h P Wuchiapingian i Paleocene Selandian 458.4 ±0.9 259.8 ±0.4 c archean P P P 61.6 i Capitanian v Darriwilian 4000 Danian 265.1 ±0.4 Middle 66.0 o 467.3 ±1.1 Guadalupian Wordian d n 268.8 ±0.5 r Dapingian 470.0 ±1.4 Hadean Maastrichtian a i Roadian O ~ 4600 72.1 ±0.2 272.3 ±0.5 Floian m Campanian r Lower 477.7 ±1.4 Units of all ranks are in the process of being defined by Global e Kungurian 283.5 ±0.6 Tremadocian Boundary Stratotype Section and Points (GSSP) for their lower 83.6 ±0.2 P boundaries, including those of the Archean and Proterozoic, long 485.4 ±1.9 Upper Santonian 86.3 ±0.5 Artinskian defined by Global Standard Stratigraphic Ages (GSSA). Charts and Cisuralian 290.1 ±0.26 Stage 10 detailed information on ratified GSSPs are available at the website Coniacian ~ 489.5 c http://www.stratigraphy.org. The URL to this chart is found below. 89.8 ±0.3 i Sakmarian 295.0 ±0.18 Furongian Jiangshanian Turonian o s ~ 494 z Asselian Paibian Numerical ages are subject to revision and do not define uni ts in c 93.9 298.9 ±0.15 u o i ~ 497 n the Phanerozoic and the Ediacaran; only GSSPs do. For boundaries o e o a Cenomanian l Gzhelian Guzhangian i in the Phanerozoic without ratified GSSPs or without constrained e Upper 303.7 ±0.1 z n 100.5 a ~ 500.5 c numerical ages, an approximate numerical age (~) is provided. a o Kasimovian 307.0 ±0.1 P n v a Series 3 Drumian s l t a Albian y ~ 504.5 s e Middle Moscovian i e s Numerical ages for all systems except Lower Pleistocene, r r u Stage 5 ~ 113.0 n 315.2 ±0.2 M Permian,Triassic, Cretaceous and Precambrian are taken from b n o C ~ 509 r e ‘A Geologic Time Scale 2012’ by Gradstein et al. (2012); Aptian Lower Bashkirian m Stage 4 e P those for the Lower Pleistocene, Permian, Triassic and Cretaceous f 323.2 ±0.4 a ~ 125.0 i Series 2 ~ 514 were provided by the relevant ICS subcommissions. n C n Upper Serpukhovian Stage 3 o Barremian a i 330.9 ±0.2 b Coloring follows the Commission for the Lower ~ 129.4 p ~ 521 r p Geological Map of the World (http://www.ccgm.org) i Hauterivian a Stage 2 ~ 132.9 s Middle Visean s C i ~ 529 Chart drafted by K.M. Cohen, S.C. Finney, P.L. Gibbard s 346.7 ±0.4 Valanginian Terreneuvian (c) International Commission on Stratigraphy, April 2016 s ~ 139.8 i Fortunian Berriasian M Lower Tournaisian To cite: Cohen, K.M., Finney, S.C., Gibbard, P.L. & Fan, J.-X. (2013; updated) ~ 145.0 358.9 ±0.4 541.0 ±1.0 The ICS International Chronostratigraphic Chart. Episodes 36: 199-204. URL: http://www.stratigraphy.org/ICSchart/ChronostratChart2016-04.pdf Aviorrhyncha magnifica Nel et al. 2013 Protoprosbole straeleni Laurentiaux, 1952 (Moscovian, Pennsylvanian; France) (Bashkirian, Pennsylvanian; Belgium) Protoprosbole Aviorrhyncha Suborder Paleorrhyncha Fam: Archescytinidae = Permopsylliidae = Lithoscytinidae = Maueriidae = Permoscytinopsidae = Uraloscytinidae = Maripsocidae = Kaltanaphididae There is a considerable diversity of Permian hemipterans, but their relationships are obscure. The supposed ancestors of the Sternorrhyncha are among the members of the suborder Paleorrhyncha. Where to search for Sternorrhyncha ancestors? Paleorrhyncha This group is a paraphyletic assemblage, ranging from the late Carboniferous (300 mya) to late Permian (270 mya). • smaller size • homonomic venation of forewings and hind wings • antennae 10-segmented with rhinaria • probably gall-making • trophic relationships with seed ferns and early gymnosperms Paleorrhyncha display several morphological specializations, e.g. variously formed ovipositors from short needle-like to very long, and forming a coil under the body. Also the placement of the rostrum base is variable. On the other hand, they show some sternorrhynchous characters, i.e. smaller size, antennae with rhinaria, and relatively simple venation. Rasnitsyn & Quicke (2002) The fossil record of the Sternorrhyncha stretches back to the Permian (270 mya); most of the fossil sites are distributed in the northern hemisphere; and only a few sites are located in the southern hemisphere. Where to search for sternorrhynchan ancestors? Sternorrhyncha • Earliest Sternorrhyncha are closely related to the Paleorrhyncha (ancestral or sister-group relationship to be revealed) • According to the fossil record, the lineage Pincombeomorpha + Aphidomorpha separated earlier than Psylloidea + Aleyrodoidea • exclusively plant sap-feeders (phloem feeders) • underwent morphological and behavioral differentiation in the Permian • have the first major radiation in the Triassic (250−200 mya) • the second in the Late Cretaceous (100−65 mya) What constitutes a sternorrhynchan insect ? First separation of Sternorrhyncha took place in the Permian (250 mya), when the infraorders Pincombeomorpha and Aphidomorpha appeared. Pincombeomorpha, with three families occur in the Permian to the Late Triassic (about 270−200 mya). Rasnitsyn & Quicke (2002) Pincombeomorpha Their characters are the forewing with the radius posterior vein (RP) originating well before the pterostigma, convex up to the nodal line and concave beyond it; nodal line crossing RP near base; median vein (M) and cubitus anterior vein (CuA) forming a short common stalk; claval vein(s) remote from posterior margin. Pincombea sp. Simulaphis shaposhnikovi Shcherbakov, 2007 Upper Permian, Australia Upper Permian, Australia During the Permian (300-250 mya), Pincombeomorpha were present in both hemispheres, but from the Triassic they are only known from the northern hemisphere. Pincombeidae (Permian-Triassic) Simulaphididae (Permian) Boreoscytidae (Permian) Evolutionary histories of each extant sternorrhynchan lineage – aphids, coccids, psyllids and whiteflies are far from complete. They are rarely fossilized because they have small and delicate bodies, and the known fossils are preserved mostly in Cretaceous and Cenozoic ambers. Next part I will briefly introduce the history of these lineages. The early evolution of aphids (Aphidomorpha) Aphidomorpha occurs in the Permian, is represented by one species.
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