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A New Suborder of Thysanura for the Carboniferous Insect Originally

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-- @ Zoological Institute, St.Petersburg, 1996 A new suborder of Thysanura for the Carboniferous insect originally described as larva of Bojophlebia, with comments on characters of the orders Thysanura and Ephemeroptera N.Ju. Kluge Kluge, N.Ju. 1996. Anew suborder of Thysanura for the Carboniferous insect originally described as larva of Bojophlebia, with comments on characters of the orders Thysanura and Ephemeroptera. ZoosysfematicaRossica, 4(1), 1995: 71-75. The fossil wingless insect originally described as lama of Bojophlebia procopi (Ephe- meroptera) belongs not to Ephemeroptera, but to Thysanura. It is named Carbotriplura kukalovae gen. et sp. n. and placed in the suborder Carbotriplurina subordo n. Here the order Thysanura is accepted in the FHrner's sense, since the widely accepted Hennig's division of Amyocerata (- Ectognatha sensu Hennig) into Monocondylia and Dicondylia is not based on actual features. Characters of Ephemeroptera and Thysanura are dis- cussed; the differences between tergaliae and paraterga and between wing buds and paranota are explained. N.Ju. Kluge, Department of Entomology, Biological Faculty, St.Petersburg State Univer- sity, Universitetskaya nab. 7, Sf.Petersburg 199034, Russia Kukalova-Peck (1985) has described 3 speci- preserved insufficiently for discussing its sys- mens of wingless insects from the Carbonifer- tematic position. No photograph of "Litho- ous of Europe and North America, which, in neura" piecko is published (there is only a her opinion, are nymphs of Ephemeroptera. drawing in the same paper, Fig. 17), thus its Two of these specimens are described as two systematic position is also obscure. The specil new species of the genus Lithoneura - L men described as the "nymph" of B. procopi piecko Kukalova-Peck, 1985 and L clayesi should be transferred to the order Thysanura. Kukalova-Peck, 1985; one specimen is at- tributed by her to the species Bojophlebia pro- Superclass HEXAPODA Latreille, 1825 copi Kukalova-Peck, 1985. The genus Litho- neura Carpenter, 1938 belongs to the family Class AMYOCERATA Remington, 1954 Syntonopteridae Handlirsch, 191 1; formerly this genus and this family were described only = Ectognatha: Hennig, 1953 (non Ectognatha Stum- from winged insects, features of their nymphs mer-Traunfels, 1891). being unknown. The monotypic genus Bo- jophlebia Kukalova-Peck, 1985 is placed in a. This taxon is characterized primarily by separate family Bojophlebiidae Kukalova- specific structure of antennae, which have Peck, 1985; the holotype of B. procopi is a muscles only in the first segment (scapus). winged insect, and the insect which is con- Since the fossilsare not adequate for recogniz- sidered to be its nymph is the only paratype of ing this character, other diagnostic characters, this species. No arguments are given to prove not so reliable, should be used. One of them is the association of these three "nymphs" with the presence, apart of paired cerci, of an un- the taxa known as winged insects, Litho- paired long paracercus, which has the same neuriidae and Bojophlebiidae respectively. secondary segmentation as cerci; such paracer- The specimen of "Lithoneura" clayesi, a cus is preserved only in Thysanura and numer- photograph of which is published (Kukalova- ous Ephemeroptera and reduced in other amy- Peck, 1985: Figs 19-21), has the structures ocerates. In all known representatives of the 72 N.Ju. Kluge: A new suborder of Thysanura ZQOSYST. ROSSICA Vd. 4 sister group of Amyocerata - the class Entog- on mandibles of various Pterygota, Collem- natha Stummer-Traunfels, 1891 - the paracer- bola, Symphyla, Chilopoda, Crustacea; hence cus is absent. Probably the presence of a pzra- such structure of mandibular condyles is plesio- cercus in Amyocerata is only a plesiomorphy, morphic, being initially present in Mandibu- but since other features are inaccessible, it may lata. Accordingly, there are no reasonable ar- be used to separate fossil Amyocerata from guments to give the Zygentoma and Microco- Entogna tha. ryphia ranks of orders. Thus the extinct Palaeo- wic taxa - Monura and CarboMpllurina subordo n. Order THYSANURA (sensu Barrier, 1904) - are also regarded only as suborders rather than orders. = Ectotrophi Grassi & Rovelli, 1890; Ectognatha Stum- mer-Traunfels, 1891; Triplura Ewing, 1942. Suborder CARBOTRIPLURINA subordo n. Here the name "Thysanura" is accepted for Diagnosis. Body not depressed laterally (in the order including the suborders Zygentoma contrast to Microcoryphia and Monura) . Par- BBrner, 1904, Microcoryphia Verhoeff, 1904, aterga of abdominal segments turned laterally Monura Sharov, 1957 and other primarily (in contrast to all other Thysanura, where wingless Amyocerata. abdominal paraterga are pressed on the lateral The name "Thysanura" has been used in sides of sternocoxa). Cerci well developed (in several different meanings.If the taxa of the contrast to Monura) . same volume are assumed identical irrespective Composition. Monotypic. of their rank, the following synonymy can be given: Thysanura Latreille, 1796 (= Aptery- Family CARBOTRIPLURIDAE fam. n. gota Lang, 1889); Thysanura sensu Lubbock, 1873 (= Cinura Packard, 1883) ; Thysanura Diagnosis and composition. The same as in sensu B'drner, 1904 (= Ectotrophi Grassi & the suborder. Rovelli, 1890) ; Thysanura sensu Crampton, 1928 (= Zygentoma BiTrner, 1904). Genus Carbotriplura gen. n. Some authors regard Zygentoma and Micro- coryphia as separate orders, assuming that Zy- Type species Carbotriplurakukalovae sp. n. gentoma is the sister group of Pterygota, and therefore combine Zygentoma and Pterygota Diagnosis and composition. The same as in into the taxon Dicondylia Hennig, 1953. The the family and suborder. mandibular structure is considered to be the basic autapomorphy of Dicondylia. Mandibles Carbotriplura kukalovae sp. n. of Zygentoma (as well as of Pterygota) are described as "dicondylous", while mandibles = Bojophlebia procopi Kukalova-Peck, 1985, partim of Microcoryphia are described as "monocon- ("nymph", non "adult"): 936, Figs 4-10. dylous". Actually, if under the tern1 "condyle" we understand a sclerotized protuberance, we Holotypus. Specimen No. P27/80, Narodni Museum, Prague, CzechRepublic (paratypus of Bojophlebiapro- have to assume that mandibles of all Zygentoma copi Kukalova-Peck, 1985). Westphalian C, base of have no such "condyles" at all. If the term whetstone horizon, "Na Stilci" quarry near Tlustice, "condyle" means a point through which the Central Bohemian Coal Basin, Bohemia. (After Kuka- axis of rotation of mandible passes (inde- lova-Peck, 1985). pendently of the presence or absence of pro- Description. See Kukalova-Peck, 1985: 936, tuberances in these points), we are to conclude Figs 4-10. that both Zygentoma and Microcoryphia have two such "condyles" and a fixed axis of rota- Discussion tion determined by them: in Microcoryphia such points of attachment are the hind pro- Tergaliae and paraterga tuberance and the end of the internal transverse ridge, while in Zygentoma both points of at- Initially Carbotriplura kukalovae has been tachment have no special morphological struc- described as mayfly nymph because its strongly tures. It is well known that two condyles in the developed lateral abdominal projections were form of sclerotized protuberances are present regarded to be tergaliae (= "tracheal gills"), ZOOSYST.ROSSICA Vol. 4 N.Ju. Kluge: A new , suborder of Thysanura 73 the appendages typical of nymphs of Ephe- num and pleuron. In the primitive state each meroptera. Actually these projections are not tergalia has the anterior and posterior costae, tergaliae, but paraterga. with branched tergalial trachea between them; The term "tergalia" (plural - "tergaliae") was in contrast to wings, where tracheae are inside introduced for the homologous paired movable veins (which are tubular costae), tergalial tra- appendages present on abdomen in Ephemero- cheae never enter into costae. ptera larvae (independently of their function), In fossils of Ephemeroptera larvae (Fig. 2), while the term "tracheal gill" was left for tergaliae are clearly distinguishable from analogous organs of any origin, which are sup- processes of other kinds, since tergaliae are plied with tracheae and are used for aquatic always well separated from their segment: apart respiration (Kluge, 1989). The term "ter- from the margins of tergaliae, the lateral mar- galiae" is derived from the term "tergum", but gins of the segment are clearly visible. it does not imply that tergaliae have originated In contrast to the tergaliae of Ephemeroptera from parts of tergum: this word means only that larvae, the lateral abdominal projections of tergaliae are in all cases attached to tergum (as Carbotriplura (Fig. 1) are connected with the well as wings do), while the origin of tergaliae remainder of the tergum all along its length, the remains vague (as well as the origin of wings). anterior margin of each projection is at the As assumed by numerous authors, tergaliae same level as the anterior margin of its segment (which are present only on abdominal seg- (if they were be tergaliae, their anterior mar- ments) may be serial homologues of wings. gins should correspond to posterior margin of Tergaliae of Ephemeroptera larvae (nymphs) their segment). Apart from the margins of have the following features (Kluge, 1989). The lateral projections, no other lateral margins of most primitive and at the same time most com- segments are visible on the published photo- mon position of tergaliae attachment is on the graphs.
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    z . G.. DAL GC Pictorial Handbook on Indian Thysanura A.K. HAZRA G.P. MANDAL Zoological Survey of India, M-Block, New Alipore, Kolkata 700 053 Edited by the Director, Zoological Survey of India, Kolkata Zoological Survey of India Kolkata CITATION Hazra, A.K. and MandaI, G.P. 2007. Pictorial Handbook on Indian Thysanura 1-43. (Published by the Director, Zool. Surv. India, Kolkata) Published : June, 2007 ISBN 978-81-8171-152-6 © Govt. of India, 2007 ALL RIGHTS RESERVED • No part of this publication may be reproduced stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise without the prior permission of the publisher. • This book is sold subject to the condition that it shall not, by way of trade, be lent, resold, hired out or otherwise disposed of without the publisher's consent, in an form of binding or cover other than that in which, it is published. • The correct price of this publication is the price printed on this page. Any revised price indicated by a rubber stamp or by a sticker or by any other means is incorrect and should be unacceptable. PRICE Indian Rs. 250.00 Foreign : $ 25; £ 20 Published at the Publication Division by the Director Zoological Survey of India, 234/4, AJe Bose Road, 2nd MSO Building, 13th floor, Nizam Palace, Kolkata 700020 and printed at MIs Image, New Delhi 110 002. FOREWORD The order Thysanura comprises of popularly known insects of (Silverfish' and 'bristle tail' Thysanurans are small, soft bodied, fishlike, scaled, and wingless insects.
  • The Phylogeny and Relationships Between the Inseet Orders

    The Phylogeny and Relationships Between the Inseet Orders

    Rev. Bio!. Trop., 8 (1): 93-123, 1960 The phylogeny and relationships between the inseet orders by Alvaro Wille'� (Received for publication May 9, 1960 ) The phylogeny and the interrelationships of the insect orders always remain a matter of great interEst to the general entomologist. The number of papers that have been published on this subject is very large. Very unfortun­ atcly, however, most of these papers are too brief, containing only a few fact3 to support the views which the author has adopted, or they deal only with one order or a group of more or less interrelated orders. The purpose of this paper is to offer a general account of the origin of the insects and the known relation­ ships between the insect orders, giving as many fact3 as possible to show these i nterrelationshi ps. As might be expected, current interpretations of the phylogeny of, and connections betwem the insect orders are not definitive, and many of our pre­ sent views may have to be changed in the future, as knowledge progresses. Although this type of studies always reveals many unsolved problems, the large amollnt of data accllmulated in the last few decades allows us to speculate on these matters, and in doing so, we may put some order and understanding into these chaotic accumulations of facts. This may explain, perhaps, the large num­ ber of papers on this subject, and will always justify further additions. Since this is not a detailed revision of al! the known views available in the literature, 1 have adopted those which the facts best seém to support.