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General System of Neoptera with Description of a New Species of Embioptera

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Russian Entomol. J. 21(4): 371384 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2012 General system of Neoptera with description of a new species of Embioptera Îáùàÿ ñèñòåìà Neoptera ñ îïèñàíèåì íîâîãî âèäà Embioptera Nikita J. Kluge Í.Þ. Êëþãå Department of Entomology, St. Petersburg State University, Universitetskaya nab., 7/9, St. Petersburg, 199034, Russia. E-mail: [email protected]. Website: http://www.insecta.bio.pu.ru Êàôåäðà ýíòîìîëîãèè, áèîëîãî-ïî÷âåííûé ôàêóëüòåò, Ñ.-Ïåòåðáóðãñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Óíèâåðñèòåòñêàÿ íàá., 7/ 8, Ñ.-Ïåòåðáóðã 199034, Ðîññèÿ. KEY WORDS: insects, systematics, phylogeny, Polyneoptera, Idioprothoraca, Notoptera, Embioptera, Rhipi- neoptera, Clothoda amazonica, new taxa. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: íàñåêîìûå, ñèñòåìàòèêà, ôèëîãåíèÿ, Polyneoptera, Idioprothoraca, Notoptera, Embioptera, Rhipineoptera, Clothoda amazonica, íîâûå òàêñîíû. ABSTRACT. Embioptera and Notoptera comprise Paraneoptera s.l.) è Metabola (= Oligoneoptera). Îá- a holophyletic taxon Idioprothoraca taxon nov., which ñóæäàþòñÿ ðàçëè÷íûå ìíåíèÿ î ñèñòåìàòè÷åñêîì is characterized by a unique structure of prothoracic ïîëîæåíèè Embioptera, Notoptera è Zoraptera. Ïî lateral walls, two-segmented cerci in the first larval èìàãî è ëè÷èíêàì èç Ïåðóâèàíñêîé Àìàçîíèè îïè- instar and some other characters. Other insect, formerly ñàí íîâûé âèä Clothoda amazonica sp.n. placed to Polyneoptera Martynov, 1923, constitute a holophyletic taxon Rhipineoptera taxon nov., which is Introduction characterized by presence of anal fan on hind wings; all representatives of Rhipineoptera taxon nov., whose anal fan is reduced, retain its vestige. In contrast to Martynov [1923, 1924, 1938] established the divi- them, Idioprothoraca taxon nov. initially have homon- sion Neoptera and divided it into three subdivisions omous wings, and their ancestors never had anal fan. Polyneoptera, Paraneoptera and Oligoneoptera. This The taxon Neoptera is divided into Idioprothoraca tax- classification became generally accepted; recently the on nov., Rhipineoptera taxon nov. and Eumetabola; the three Martynovs subdivisions are widely recognized taxon Eumetabola is divided into Parametabola (= Para- either as subdivisions, or cohorts [e.g., Rohdendorf, neoptera s.l.) and Metabola (= Oligoneoptera). Various 1962], or supercohorts [e.g., Arillo & Engel, 2006], or opinions about systematic position of Embioptera, No- infraclasses [e.g., Kukalova-Peck, 1973], or rankless toptera and Zoraptera are discussed. New species Clotho- taxa. Among them, Oligoneoptera is a holophyletic da amazonica sp.n. is described basing on male ima- taxon of insects with complete metamorphosis, whose goes and nymphs from Peruvian Amazonia. oldest mane is Metabola (see below). The taxon Parane- optera has a more monosemantic name Parametabola; I ÐÅÇÞÌÅ. Embioptera è Notoptera îáðàçóþò ãîëî- regard it to be holophyletic, while another opinion ôèëåòè÷åñêèé òàêñîí Idioprothoraca taxon nov., êîòî- exists (see discussion on Zoraptera below). Hennig ðûé õàðàêòåðèçóåòñÿ óíèêàëüíûì ñòðîåíèåì ëàòå- [1953] assumed that Metabola and Parametabola com- ðàëüíûõ ñòåíîê ïåðåäíåãðóäè, äâó÷ëåíèêîâûìè öåð- prise a holophyletic taxon Eumetabola, that seems reli- êàìè ó ëè÷èíêè ïåðâîãî âîçðàñòà è íåêîòîðûìè äðó- able (see below). The taxon Polyneoptera was original- ãèìè ïðèçíàêàìè. Ïðî÷èå íàñåêîìûå, ðàíåå ïîìå- ly characterized by a single apomorphic character ùàâøèåñÿ â Polyneoptera Martynov, 1923, îáðàçóþò presence of anal fan on hind wing. Some authors ex- ãîëîôèëåòè÷åñêèé òàêñîí Rhipineoptera taxon nov., press doubt that this apomorphy testifies about holo- êîòîðûé õàðàêòåðèçóåòñÿ íàëè÷èåì àíàëüíîãî âååðà phyly of Polyneoptera, so they prefer to call this group íà çàäíèõ êðûëüÿõ; ó âñåõ ïðåäñòàâèòåëåé Rhipi- lower Neoptera [e.g., Kristensen, 1995], but no other neoptera taxon nov., ó êîòîðûõ àíàëüíûé âååð ðåäó- classification has been suggested. Among taxa included öèðîâàí, ñîõðàíÿþòñÿ åãî âåñòèãèè.  îòëè÷èå îò to Polyneoptera, there is an order Embioptera; Mar- íèõ, ó Idioprothoraca taxon nov. êðûëüÿ èñõîäíî ãî- tynov mentioned this order as a member of Polyneoptera ìîíîìíûå, è èõ ïðåäêè íèêîãäà íå èìåëè àíàëüíîãî in all his publications about general insect systematics, âååðà. Òàêñîí Neoptera äåëèòñÿ íà Idioprothoraca including even the first one [Martynov, 1923], in which taxon nov., Rhipineoptera taxon nov. è Eumetabola; the list of taxa included was very incomplete. Unlike òàêñîí Eumetabola äåëèòñÿ íà Parametabola (= other members of Polyneoptera, Embioptera have nei- 372 N.J. Kluge ther anal fan, nor its vestige. Thus, the taxon Poly- margin of pseudopleurite. As pleurites of prothorax are di- neoptera has no any autapomorphies and even has no minished and located far from anterior margin of prothorax, formal diagnosis. In this paper I give a modified classi- cervical sclerites, being articulated with them, are elongated fication, where insects formerly placed to an artificial and stretch from pleurites to the head, being located ventrad taxon Polyneoptera, form two independent taxa Idio- of pseudopleurites. Each cervical sclerite is divided into two sclerites anterior and posterior ones; the posterior cervical prothoraca taxon nov. and Rhipineoptera taxon nov. sclerite is transverse, short and wide; the anterior cervical Names of high-rank insect taxa are given according sclerite is longitudinal, long, narrowing anteriorly. Thus, in to my previous paper [Kluge, 2010a], in agreement with Idioprothoraca neck is short, but neck sclerites are long. the principles of circumscriptional nomenclature, which Structure of prothoracic lateral wall strongly differs from constituted a part of the dual nomenclature system. structure of meso- and metathoracic lateral walls [see (3)]. Some figures used in this paper, were made from The presence of pseudopleurite is a unique feature of specimens of Embioptera belonging to an undescribed Idioprothoraca; apodeme, formed by dorsal portion of pleur- species. Here this species is described as Clothoda ama- ite (cryptopleurite), besides Idioprothoraca, is found also in zonica sp.n. Type material of this species is deposited in Saltatoria and Coleoptera. the Zoological Institute of Russian Academy of Sciences. (2) Cerci are 2-segmented at least in first-instar larva. In Embioptera cerci remain to be 2-segmented in all instars I. Proposed classification and characteristics of taxa (Figs 6, 8). In Notoptera cerci are 2-semented in the first instar only [Fig. 3; Nagashima, 1991: Fig. 3; Uchifune & 1. NEOPTERA Martynov, 1923 Machida, 2005: Figs 4AC, 11]; at each molt number of Circumscriptional synonymy: segments increases thanks to division of the most proximal = NEOPTERYGOTA Crampton, 1924. segment, so that in imago cercus has 510 segments. In other Typified name in basic format: Scarabaeus/fg (sine Libellula; insects cerci either consist of indeterminate number of seg- incl. Embia, Forficula). ments beginning from the first instar (e.g., many Plecoptera, Autapomorphy. Wings have got ability to fold thanks to Pandictyoptera), or are one-segmented in all instars (e.g., mobility of the 3rd axillary sclerite; when the 3rd axillary Saltatoria, Spectra), or are multisegmented in larva and one- sclerite turns over, the wing turns by its apex backward, segmented in imago (e.g., some Plecoptera and some Der- keeping its dorsal side dorsally. matoptera); only in Tridactylidae cerci are 2-segmented. Classification. The taxon Neoptera is divided into Idio- Probable apomorphies of Idioprothoraca: prothoraca, Rhipineoptera and Eumetabola. (3) Mesothorax and metathorax have similar structure different from structure of prothorax [see (1)]. Pleurites are 1.1. IDIOPROTHORACA taxon nov. much larger than that of prothorax, and strongly inclined, so Typified name in basic format: Embia/fg (incl. Grylloblatta). that pleural suture stretches longitudinally nearly from ante- ETYMOLOGY. éäéïò (special) + prothorax; due to au- rior margin of the tergum to posterior margin of the segment. tapomorphy in prothoracic structure [see character (1)]. In accordance with this, in winged forms (male Embioptera CHARACTERISTICS. Autapomorphies of Idioprothoraca: only) wings are attached near anterior margin of the segment (1) Prothorax is modified as the following: pleurites are (Fig. 7). In larvae of these winged forms protopteron begins strongly diminished, and tergite is elongate in front of the from anterior margin of tergum, and its epipleuron forms the pleurites; side walls of the prothorax in front of pleurites are whole lateral margin of the tergum (Fig. 5). In wingless forms formed by a pair of pseudopleurites newly formed scler- (females of Embioptera and both sexes of Notoptera) the ites of a characteristic shape; dorsal part of the pleurite is same epipleura of protoptera are present, being developed dipped into the body, forming a sclerotized apodeme (Figs 1 both in larvae and imagoes; they represent lamellate lateral 2, 4, 7). Pseudopleurite forms a part of lateral wall of protho- margins of terga, which are bent down and cover dorsal areas rax, but probably have originated from a lateral portion of of pleurites (Figs 1, 4) (the name Notoptera is connected tergite. In Embioptera, pseudopleurite is flat or somewhat with this feature). convex and exposed laterally or laterally-dorsally; it can be Possibly, the common ancestor of Idioprothoraca had wings separated from notum (i.e., dorsal part of tergite) by a mem- in males only, while its females were wingless (as recent branous stripe or two membranous
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  • Changes to the Fossil Record of Insects Through Fifteen Years of Discovery

    Changes to the Fossil Record of Insects Through Fifteen Years of Discovery

    This is a repository copy of Changes to the Fossil Record of Insects through Fifteen Years of Discovery. White Rose Research Online URL for this paper: https://eprints.whiterose.ac.uk/88391/ Version: Published Version Article: Nicholson, David Blair, Mayhew, Peter John orcid.org/0000-0002-7346-6560 and Ross, Andrew J (2015) Changes to the Fossil Record of Insects through Fifteen Years of Discovery. PLosOne. e0128554. https://doi.org/10.1371/journal.pone.0128554 Reuse Items deposited in White Rose Research Online are protected by copyright, with all rights reserved unless indicated otherwise. They may be downloaded and/or printed for private study, or other acts as permitted by national copyright laws. The publisher or other rights holders may allow further reproduction and re-use of the full text version. This is indicated by the licence information on the White Rose Research Online record for the item. Takedown If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request. [email protected] https://eprints.whiterose.ac.uk/ RESEARCH ARTICLE Changes to the Fossil Record of Insects through Fifteen Years of Discovery David B. Nicholson1,2¤*, Peter J. Mayhew1, Andrew J. Ross2 1 Department of Biology, University of York, York, United Kingdom, 2 Department of Natural Sciences, National Museum of Scotland, Edinburgh, United Kingdom ¤ Current address: Department of Earth Sciences, The Natural History Museum, London, United Kingdom * [email protected] Abstract The first and last occurrences of hexapod families in the fossil record are compiled from publications up to end-2009.