Evolution of the Beetle Hind Wing, With Special Reference to Folding (Insecta, Coleoptera)

337 Pages·2009·6.34 MB·English

by  Dmitri N. Fedorenko

Dr. Dmitri Fedorenko, born 1962, WD EVOLUTION
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is Senior Scientist at the Institute T
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of Ecology and Evolution, r
Russian Academy of Sciences, Moscow. SPi N OF THE BEETLE HIND WING,
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His main interests lie in the taxonomy, CF
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ecology, geography and evolution of the Ad WITH SPECIAL REFERENCE
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beetles, the ground-beetles in particular. r
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He is the author of more than Ek
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40 scientific papers, including
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the monograph “Reclassification
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of world Dyschiriini, with a revision N
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of the Palearctic fauna E
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(Coleoptera, Carabidae)”, TL
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Moscow–Sofia–St. Petersburg: T
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Pensoft Publishers, 1996. OO
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Dmitri N. Fedorenko
The present monograph is a highly original and thorough IO
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attempt at revising the wing structure of the beetles, with G
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special emphasis placed not only on the venation patterns N
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observed, but also on folding. Combined, all of these patterns EB
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are critically re-evaluated to provide new, highly unorthodox TE
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insights in beetle evolution. The work is also abundantly ,L

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illustrated by original drawings showing all the necessary O H
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details of beetle wing structure, including shape, venation, EN
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sclerotization and folding patterns. The present monograph P
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is indispensable for students in beetle taxonomy, evolution EI
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and palaeontology. AG
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ISBN 978-954-642-494-5 (PB)
ISBN 978-954-642-495-2 (e-book)
EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING 1
EVOLUTION OF THE BEETLE HIND WING,
WITH SPECIAL REFERENCE TO FOLDING
(INSECTA, COLEOPTERA)
2 EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING
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EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING 3
A. N. S I E E
EVERTZOV NSTITUTE OF COLOGY AND VOLUTION
R A S
USSIAN CADEMY OF CIENCES
EVOLUTION
OF THE BEETLE HIND WING,
WITH SPECIAL
REFERENCE TO FOLDING
(INSECTA, COLEOPTERA)
by
DMITRI N. FEDORENKO
Edited by
Sergei I. Golovatch
Sofia –Moscow
2009
4 EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING
EVOLUTION OF THE BEETLE HIND WING,
WITH SPECIAL REFERENCE TO FOLDING
(INSECTA, COLEOPTERA)
by
Dmitri N. Fedorenko
Edited by
Sergei I. Golovatch
First published 2009
ISBN: 978-954-642-494-5 (paperback)
ISBN: 978-954-642-495-2 (e-book)
© PENSOFT Publishers
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system or transmitted in any form by any means,
electronic, mechanical, photocopying, recording or otherwise,
without the prior written permission of the copyright owner.
Pensoft Publishers
Geo Milev Str. 13a, Sofia 1111, Bulgaria
[email protected]
www.pensoft.net
Printed in Bulgaria, July 2009
EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING 5
TABLE OF CONTENTS
7
Introduction
9
Material, methods and terminology
17
Structure and operation of insect wings: some backgrounds 
17
Th e insect wing 
21
Th e hind wing of Coleoptera
27
Wing structure: basic elements and their genesis through evolution
27
Fore wing, or elytron
29
Wing venation and elytra-body interlocking
33
Hind wing
35
Wing shape
38
Th e articular region 
49
Fold- and fl exion-lines
51
Hind wing venation
74
Secondary supporting structures
78
Folding pattern
87
Evolution of the hind wing in Coleoptera
87
Th e main folding patterns (folding types)
102
Th e folding apparatus
110
Th e supporting system
119
Th e wing as a whole
120
Morphological and morphofunctional types
132
Th e main evolutionary trends and functional wing types
135
Size evolution of the imago
139
Conclusion
143
Hind wing structure and evolution in particular beetle groups
143
Adephaga
155
Myxophaga
157
Scirtoidea
159
Staphyliniformia s.l.
162
Hydrophiloidea s.l.
6 EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING
164
Scarabaeoidea
171
Staphylinoidea
174
Elateriformia s.str.
178
Elateroidea
184
Byrrhoidea
185
Buprestoidea + Dascilloidea
187
Th e cucujiform wing
188
Derodontoidea
189
Bostrichoidea
191
Lymexiloidea
192
Tenebrionoidea
198
Cucujoidea
201
Chrysomeloidea, Curculionoidea and Cleroidea
202
Cleroidea
204
Phytophaga
204
Chrysomeloidea
205
Curculionoidea
Morphological evolution of the beetle hind wing and the
209
phylogeny of Coleoptera
213
Relationships of individual beetle taxa based on hind wing structure
213
Adephaga
224
Staphyliniformia
229
Elateriformia
236
Bostrichoidea
237
Tenebrionoidea
243
Cucujoidea 
248
Chrysomeloidea
248
Curculionoidea
250
References
263
Appendix 1
323
Appendix 2
333
Appendix 3
EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING 7
INTRODUCTION
Th e beetles, Coleoptera, or Scarabaeida, is the most diverse insect order com-
prising about 360,000 extant species. Th ese are arranged into four suborders, a
few series, 16 superfamilies and ca 160–170 families in the most recent classi-
fi cations (e.g. Lawrence & Newton, 1995). Among holometabolan insects (En-
dopterygota), the Coleoptera is prominent because its origin seems to have been
a direct result of strong transformations of its fl ight apparatus. More specifi cally,
archetypal beetle features, especially the adult external structure transformed
into a durable protective construction, resulted immediately from a strongly
increased defensive function of the fore wings. Having transformed into rigid
elytra, these reliably prevented the hind wings in repose and the hindbody terga
from damage or some other troubles when the imago penetrated into any liq-
uid, closed or running substrate, or, on the contrary, left a confi ned space after
preimaginal development.
Formation of such a structural plan of the adult body brought about pro-
found change in the entire fl ight apparatus. Functional posteromotorism, the
hind wings longer than the elytra and thence reducible in length when not in
use were the main sequels to this process. Th e great variety of body shapes and
sizes, together with the immense taxonomic diversity of Coleoptera, contributed
much to the diversity of hind wing venation and folding patterns. Neverthe-
less, the understanding of the general trends that defi ned this diversity is far
from complete. Many studies have been restricted to hind wing venation in
separate beetle families or, more seldom, superfamilies. Comparative analyses
of venational characters across the order are singular. Some of them are out
of date while some others seem superfi cial. In particular, Ganglbauer’s (1892)
well-known venational typology has been shown to be largely artifi cial, with
its carabidiform, cantharidiform and staphylinidiform venation types often being
only stages of successive veinal reductions (Ponomarenko, 1972). A compara-
tively recent typology introduced by Wallace & Fox (1980) has not been widely
adopted. Special emphasis was put on folding patterns by Forbes (1926), but
with no or minimum relation to wing venation. As a result, Forbes’ typology
proved to largely be functional. Nonetheless, the placement of a few families
in the Adephaga, for which a separate suborder, Myxophaga, was afterwards
8 EVOLUTION OF THE BEETLE HIND WING, WITH SPECIAL REFERENCE TO FOLDING
erected (Crowson, 1955), as well as the incorporation of the Hydraenidae into
the Staphylinoidea, and the Haplogastra shown to be a natural group became
greatly appreciated novelties of his classifi cation based on this typology (Law-
rence et al., 1995b).
Th us, phylogenetic reconstructions and higher level classifi cations of the
order, proposed to date from analyses of wing structure, have been supported by
separate characters regarded as set elements, not parts of the whole, these be-
ing interrelated and thence interdependent in their changes through evolution.
Such an approach obscures the intrinsic trends of evolution both of the wing as
a whole and of its integral parts, forbidding the separation of synapomorphies
from homoplasies. Hence, it leads to warped results and is less effi cient. Th e
objective of this work is to fi ll in this gap.
MATERIAL, METHODS AND TERMINOLOGY 9
MATERIAL, METHODS AND TERMINOLOGY
Hind wing structure, namely venation, articular region, folding pattern, trache-
ation of the main veinal stems, sclerotization patches, fl exion lines, and mesore-
lief of the wing membrane were studied in members of Recent beetle families
listed in Appendix 2, as well as in individual representatives of Ephemeroptera,
Orthoptera, Dermaptera, Dictyoptera, Plecoptera, Megaloptera, Neuroptera,
Trichoptera, Lepidoptera, Mecoptera, Diptera and Hymenoptera. Some of
these are depicted (Figs 8, 9, 12–14, 22–24).
Unfolded wings were mounted on glass slides with a transparent adhesive
and then examined under the stereoscope at magnifi cations of 8× to 56×.
All convex (mountain) and concave (valley) folds were identifi ed in isolated,
non-fi xed, mostly dry wings; these were examined fi rst folded and then
stepwise unfolded in water or glycerol. Mesorelief peculiarities of the wing
membrane were explored in a drying drop of water. Th e left and right wings
were largely examined using more than one specimen, usually 2–10, of each
species listed. Most of the folding patterns thus obtained were checked on
in paper models.
Th e following three parameters were measured: the length of the body
(BL), the length of the wing (WL) and the length of the apical membrane
(ML) (Appendix 3), with further analyses of the indices WL/BL (%) and ML/
WL (%). WL was measured in a fully unfolded wing, from its apex to the pos-
terobasal margin or articular region in the case of a strongly reduced jugum or
clavus plus jugum. ML was measured from wing tip to the anterior tip of the
apical hinge or to the distal costal pivot (Polyphaga), or to the proximal costal
pivot (Archostemata, Adephaga). In the text, the term long(er) wing or long(er)
apical membrane corresponds to a high(er) value of WL/BL or ML/WL ratio,
respectively, unless otherwise stated.
Information on the structure of fore and hind wings of extinct Coleoptera
was predominantly extracted from the literature (Ponomarenko, 1969, 1972,
1975; Nikritin, 1977; Nikolajev, 1995; Krell, 2006). Only the hind wing imprint
of an Upper Permian insect was examined, probably that of a beetle labelled
“PIn AS USSR, № 3286/4±, Udmurt ASSR, right bank of Kama River, di-
viding range of Rassokha and Kasyanovka rivers, 1.5 km NE of Chepanikha,