History of Insects

523 Pages·2002·130.239 MB·English

by  N. V. Belayeva, V. A. Blagoderov, V. Yu. Dmitriev, K. Yu. Eskov, A. V. Gorokhov, V. D. Ivanov, N. Yu. Kluge, M. V. Kozlov, E. D. Lukashevich, M. B. Mostovski, V. G. Novokshonov, A. G. Ponomarenko, Yu. A. Popov, L. N. Pritykina, A. P. Rasnitsyn, D. E. Shcherbakov, N. D. Sinitshenkova, S. Yu. Storozhenko, I. D. Sukatsheva, V. N. Vishniakova, Peter Vršanský, V. V. Zherikhin (auth.), Alexandr P. Rasnitsyn, Donald L.J. Quicke (eds.)

HISTORY OF INSECTS
HISTORY OF INSECTS
by
N.V. Belayeva, V.A. Blagoderov, V.Yu. Dmitriev, K.Yu. Eskov, A.V. Gorokhov,
V.D. Ivanov, N.Yu. Kluge, M.V. Kozlov, E.D. Lukashevich, M.B. Mostovski,
V.G. Novokshonov, A.G. Ponomarenko, Yu.A. Popov, L.N. Pritykina, A.P. Rasnitsyn,
D.E. Shcherbakov, N.D. Sinitshenkova, S.Yu. Storozhenko, I.D. Sukatsheva,
V.N. Vishniakova, Peter Vršanský and †V.V. Zherikhin
Edited by
ALEXANDR P. RASNITSYN AND DONALD L.J. QUICKE
KLUWER ACADEMIC PUBLISHERS
NEW YORK, BOSTON, DORDRECHT, LONDON, MOSCOW
eBookISBN: 0-306-47577-4
Print ISBN: 1-4020-0026-X
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This book is dedicated to the three people
most responsible for the present state of palaeoentomology,
Andrey Vassilievich Martynov,
Boris Borisovich Rohdendorf and
Frank Morton Carpenter
We also extend our dedication to
Vladimir Vassilievich Zherikhin, whose untimely
death in December 2001,
was a great loss to
palaeoentomology
Contents
Preface xi 1.4.3.3. Subaerial palaeoenvironments 57
Acknowledgements xi 1.4.3.4. Selectivity of the rock record 60
1.4.3.5. Insect microfossils and chemofossils 60
Contributors xii
1.4.4. Insects as contaminants in fossil assemblages 61
1.4.5. Insect activities as a taphonomic factor 62
1. INTRODUCTION TO PALAEOENTOMOLOGY 1
1.1. Scope and approach 1
A.P. Rasnitsyn 2. CLASS INSECTA Linné, 1758. THE INSECTS 65
1.1.1. Coverage 1 A.P.Rasnitsyn
1.1.2.Phylogenetic approach 1 2.1. Subclass Lepismatona Latreille, 1804. 69
1.1.3.Nomenclature 7 The wingless insects
1.2. Special features of the study of fossil insects 8 2.1.1. Order Machilida Grassi, 1888. The bristletails 70
A.P.Rasnitsyn 2.1.2. OrderLepismatida Latreille, 1804. Thesilverfish 73
1.3. Concise history of palaeoentomology 12 2.2. Subclass ScarabaeonaLaicharting, 1781.
A.P.Rasnitsyn The winged insects 75
1.4. Pattern of insect burial andconservation 17 2.2.0.1. Order Paoliida Handlirsch, 1906 83
†V.V.Zherikhin 2.2.1. Infraclass Scarabaeones Laicharting, 1781 84
1.4.1. General 17 2.2.1.1. CohorsLibelluliformes Laicharting, 1781 85
1.4.2. Insect taphonomy 18 2.2.1.1.1. Superorder Ephemeridea Latreille, 1810.
1.4.2.1. Direct burial in sedimentary deposits 18 The mayflies 86
1.4.2.1.1. Autotaphonomical factors 19 2.2.1.1.1.1. OrderTriplosobida Handlirsch, 1906 87
1.4.2.1.2. Ecological factors 24 2.2.1.1.1.2. Order Syntonopterida Handlirsch, 1911 88
1.4.2.1.2.1. Ecological factors affecting organisms in their 2.2.1.1.1.3. OrderEphemerida Latreille, 1810.
life-time 25 The true mayflies 89
1.4.2.1.2.2. Mortality factors 27 N.Yu. Kluge and N.D. Sinitshenkova
1.4.2.1.2.3. Post-mortem ecological factors 29 2.2.1.1.2. Superorder Libellulidea Laicharting, 1781.
1.4.2.1.3. Taphotopical factors 31 OrderOdonata Fabricius, 1792. The dragonflies 97
1.4.2.1.4. Postburial factors 36 A.P. Rasnitsyn and L.N. Pritykina
1.4.2.1.5. Technical factors 39 2.2.1.2. Cohors Cimiciformes Laicharting, 1781 104
1.4.2.2. Indirect burial in fossil containers 40 A.P. Rasnitsyn
1.4.2.2.1. Fossil resins 40 2.2.1.2.1. Superorder Caloneuridea Handlirsch, 1906 105
1.4.2.2.1.1. Autotaphonomical factors 41 2.2.1.2.1.1. Order Blattinopseida Bolton, 1925 106
1.4.2.2.1.2. Ecological factors 43 2.2.1.2.1.2. OrderCaloneurida Handlirsch, 1906 106
1.4.2.2.1.3. Taphotopical factors 43 2.2.1.2.1.3. Order Zorotypida Silvestri, 1913 109
1.4.2.2.1.4. Postburial factors 48 2.2.1.2.2. SuperorderHypoperlidea Martynov, 1928.
1.4.2.2.1.5. Technical factors 50 OrderHypoperlida Martynov, 1928 111
1.4.2.2.2. Other primary fossil containers 50 2.2.1.2.3. Superorder Dictyoneuridea Handlirsch, 1906 115
1.4.2.2.3. Secondary fossil containers 52 N.D.Sinitshenkova
1.4.3. Products of the taphonomical process: insect fossils 2.2.1.2.3.1. OrderDictyoneurida Handlirsch, 1906 116
and ichnofossils in different palaeoenvironments 2.2.1.2.3.2. Order Mischopterida Handlirsch, 1906 120
and modes of their preservation 53 2.2.1.2.3.3. Order Diaphanopterida Handlirsch, 1906 123
1.4.3.1. Marine deposits 53 2.2.1.2.4. Superorder Psocidea Leach 1815 125
1.4.3.2. Non-marine subaquatic palaeoenvironments 54 A.P.Rasnitsyn
1.4.3.2.1. Lacustrine deposits 54 2.2.1.2.4.1. Order Psocida Leach, 1815. The booklice 128
1.4.3.2.2. Swamp, marsh and other wetland deposits 56 2.2.1.2.4.2. OrderPediculida Leach, 1815. The lice 131
1.4.3.2.3. Fluvial deposits 56 2.2.1.2.4.3. OrderThripida Fallen, 1914. 133
1.4.3.2.4. Spring deposits 57 †V.V. Zherikhin
vii
CONTENTS
2.2.1.2.5. Superorder Cimicidea Laicharting, 1781 2.2.2.2.4. Order Embiida Burmeister, 1835.
Order Hemiptera Linné, 1758. The bugs, The webspinners 291
cicadas, plantlice, scale insects, etc. 143 A.P.Rasnitsyn
D.E. Shcherbakov and Yu.A. Popov 2.2.2.3. Superorder Gryllidea Laicharting, 1781 293
2.2.1.3. Cohors Scarabaeiformes Laicharting, 1781 A.V.Gorochov and A.P. Rasnitsyn
The holometabolans 157 2.2.2.3.1. OrderOrthoptera Olivier, 1789. The orthopterans 294
A.P. Rasnitsyn 2.2.2.3.2. OrderPhasmatida Leach, 1915. The stick insects 301
2.2.1.3.0.1. Order Stylopida Stephens, 1829 159 2.2.2.3.3. Order Mesotitanida Tillyard, 1925 302
2.2.1.3.1. Superorder Palaeomanteidea Handlirsch, 1906. 2.3. Insect trace fossils 303
OrderPalaeomanteidaHandlirsch, 1906 161 †V.V.Zherikhin
2.2.1.3.2. Superorder Scarabaeidea Laicharting, 1781. 2.3.1. Introductory remarks 303
Order Coleoptera Linné, 1758. The beetles 164 2.3.2. Locomotion and resting traces 307
A.G. Ponomarenko 2.3.3. Shelters and burrows 308
2.2.1.3.3. SuperorderMyrmeleontidea Latreille, 1802 176 2.3.4. Oviposition traces 316
A.G. Ponomarenko 2.3.5. Feeding traces 318
2.2.1.3.3.1. OrderRaphidiida Latreille, 1810 177 2.3.6. Coprolites 324
2.2.1.3.3.2. Order Corydalida Leach, 1815 180
2.2.1.3.3.3. OrderNeuropteraLinné, 1758 183
2.2.1.3.3.4. OrderJurinida M. Zalessky, 1928 189 3. GENERAL FEATURES OF INSECT HISTORY 325
A.P. Rasnitsyn 3.1. Dynamics of insect taxonomic diversity 325
2.2.1.3.4. Superorder Papilionidea Laicharting, 1781 192 V.Yu. Dmitriev and A.G. Ponomarenko
A.P. Rasnitsyn 3.2. Ecological history of terrestrial insects 331
2.2.1.3.4.1. OrderPanorpida Latreille, 1802. †V.V.Zherikhin
The scorpionflies 194 3.2.1. Introduction 331
V.G. Novokshonov 3.2.2. Palaeozoic 332
2.2.1.3.4.2. Order Trichoptera Kirby, 1813. 3.2.2.1. Environments of insect origin and earlyevolution 332
The caddisflies 199 3.2.2.2. Middle carboniferous to permian 335
V.D. Ivanov and I.D. Sukatsheva 3.2.2.2.1. General features of palaeozoic insects 335
2.2.1.3.4.3. Order Lepidoptera Linné, 1758. 3.2.2.2.2. Herbivores 338
The butterflies and moths 220 3.2.2.2.3. Detritivores and fungivores 343
M.V. Kozlov, V.D. Ivanov and A.P. Rasnitsyn 3.2.2.2.4. Predators 345
2.2.1.3.4.4. Order Diptera Linné, 1758. The true flies 227 3.2.2.2.5. Some ecosystem-level phenomena 347
V.A. Blagoderov, E.D. Lukashevich 3.2.2.3. The palaeozoic/mesozoic transition 348
and M.B. Mostovski 3.2.3. Mesozoic 351
2.2.1.3.4.5. OrderPulicida Billbergh, 1820. 3.2.3.1. Triassic – early cretaceous 351
The fleas 240 3.2.3.1.1. General features of mesozoic insects
A.P. Rasnitsyn (beforemid-cretaceous) 351
2.2.1.3.5. Superorder Vespidea Laicharting, 1781. 3.2.3.1.2. Herbivores 352
Order Hymenoptera Linné, 1758 242 3.2.3.1.2.1. Anthophily and the evolution of entomophily 353
A.P. Rasnitsyn 3.2.3.1.2.2. Other forms of herbivory 358
2.2.2. Infraclass Gryllones Laicharting, 1781. 3.2.3.1.3. Detriti- and fungivores 363
The grylloneans 254 3.2.3.1.4. Predators and parasites 365
A.P. Rasnitsyn 3.2.3.1.5. Origin of sociality 367
2.2.2.0.1. Order Eoblattida Handlirsch, 1906 256 3.2.3.1.6. Some ecosystem-level phenomena 367
2.2.2.1. Superorder Blattidea Latreille, 1810 260 3.2.3.2. The late cretaceous and the mesozoic/cainozoic
2.2.2.1.1. Order Blattida Latreille, 1810. The cockroaches 263 transition 367
Peter Vršanský, V.N. Vishniakova and A.P. Rasnitsyn 3.2.3.2.1. Generalfeatures of late cretaceous insects 367
2.2.2.1.2. OrderTermitida Latreille, 1802. The termites 270 3.2.3.2.2. Herbivores 369
N. V. Belayeva 3.2.3.2.3. Detriti-andfungivores 371
2.2.2.1.3. OrderMantida Latreille, 1802. The mantises 273 3.2.3.2.4. Predators and parasites 372
†V.V. Zherikhin 3.2.3.2.5. Pattern and models of the mid-cretaceous events 373
2.2.2.2. SuperorderPerlideaLatreille, 1802 276 3.2.4. Cainozoic 374
A.P. Rasnitsyn 3.2.4.1. General features of cainozoic insects 374
2.2.2.2.1. Order Grylloblattida Walker, 1914 278 3.2.4.2. Herbivores 376
S.Yu. Storozhenko 3.2.4.3. Detriti- and fungivores 382
2.2.2.2.2. OrderPerlidaLatreille, 1810. The stoneflies 281 3.2.4.4. Predators and parasites 384
N.D. Sinitshenkova 3.2.4.5. Insect sociality and someecosystem
2.2.2.2.3. OrderForficulida Latreille, 1810. level phenomena 385
The earwigs and protelytropterans 288 3.2.4.6. Origin of principal types of modern terrestrial
D.E. Shcherbakov ecosystems 386
viii
CONTENTS
3.3. Ecological history of the aquatic insects 388 3.4.3. Carboniferous stage of insect evolution 428
N.D. Sinitshenkova 3.4.4. Permian stage of insect evolution 429
3.3.1. Introductory notes 388 3.4.5. Triassic stage of insect evolution 430
3.3.2. Carboniferous 389 3.4.6. Jurassic stage of insect evolution 431
3.3.3. Permian 390 3.4.7. Cretaceous stage of insect evolution 431
3.3.4. Triassic 391 3.4.8. Cainozoic stage of insect evolution 433
3.3.4.1. Overview of taxa of aquatic insects 391 3.4.9. Problem of the so-called “Gondwanan”
3.3.4.2. Aquatic insect assemblages and their typology 393 Ranges of the recent taxa 433
3.3.5. Jurassic 394 3.4.10. Somegeneral phylogenetic patterns 434
3.3.5.1. Overview of taxa of aquatic insects 394
3.3.5.2. Aquatic insect assemblages and their typology 398 4. APPENDIX: SELECTED INSECT FOSSIL SITES 437
3.3.6. Cretaceous 400 4.1. Impression fossils 437
3.3.6.1. Overview of taxa of aquatic insects 401 A.P. Rasnitsyn and†V.V. Zherikhin
3.3.6.2. Aquatic insect assemblages and their typology 404 4.2. Fossil resins 444
3.3.7. Cainozoic 417 K.Yu. Eskov
†V.V. Zherikhin and N.D. Sinitshenkova
3.3.7.1. Overview of taxa of aquatic insects 417 5. REFERENCES 447
3.3.7.2. Aquatic insect assemblages and their typology 418
3.3.7.3. Models of cretaceous and cainozoic evolution of INDEXES
aquatic insect assemblages 425 General index 495
3.4. Geographical history of insects 427 Index to taxon names 498
K.Yu. Eskov Index to straton names 511
3.4.1. Introductory remarks 427 Index to geographic names 513
3.4.2. Devonian stage of insect evolution 427
ix
Preface
Insects are not dinosaurs – and they probably pose us more strange ACKNOWLEDGEMENTS
puzzles and unexpected questions. A million extant species, that is sev-
For A.P. Rasnitsyn, preparing of various parts of this book was sup-
eral times more than all other living taxa together, is still a very con-
ported in part by grants: by the International Science Foundation, by
servative estimate, and their real number is for sure many times more.
the Leverhulme Trust to D.L.J. Quicke and M.G. Fitton; by the Royal
They are incomparably diverse in terms of their size, structure and way
Society Joint Project with the FSU to APR and E.A. Jarzembowski; by
of life – and yet they are all small – by our standard at least – why? And
ESF Project ‘Fossil Insects Network’; by RFFI grants 95-04-11105,
they practically ignore the cradle of life, the sea – again, why? Of
98-04-48518; by the Smithsonian Institution and California Academy
course, some survive and even reproduce in salt water, but nevertheless
of Sciences; and by various help, including sharing unpublished
very few of them are specialised for marine life. Some insects have
information, rendering material, advice, publications, shelter and work-
developed highly elaborate forms of sociality, far surpassing all
ing facilities from many friends and colleagues, of which APR would
achievements reached by vertebrates (including ourselves), at least in
like to mention specially, besides the co-editor and all co-authors of
terms of interdependence of individuals. Once again, why have they
the present book, also Dr. H.H. Basibuyuk, Dr J.M. Carpenter,
developed this way of life? The history of insects is also full of unex-
Mr. J. Cooper, Mr R. Coram, Dr. H. Dathe, Dr. E.A. Jarzembowski,
pected discoveries as well as of painful gaps, though these omissions
Dr. C.C. Labandeira, Mr. D. Kohls, Dr. J. Kukalov-Peck, Dr. X. Martìnes-
are not overwhelming nor necessarily senseless. There is a special
Delclòs, Dr. L. Masner, Dr. A. Nel, Mr. L. Pribyl, Dr W.J. Pulawski, Dr.
branch of palaeontology calledtaphonomy whose aim is to learn about
A.J. Ross, Dr. J.M. Rowland, Dr. W. Zessin.
the burial pattern of the past organisms: what governs their chance of
For V.D. Ivanov, this study was supported by the RFFI grants N
becoming fossilised, and how this chance depends on features of both
99-04-49564, 00-15-97934, by the Federal program ‘Universities of
the organisms themselves and on their environments. Depending on
Russia’ (project N 3917), and by project BR-7 of the program
how deep the taphonomical background in a particular research field,
‘Bioraznoobrazie’.
it is even possible to glean information from the very gaps in the fossil
N.Yu Kluge’s contribution was supported by the Federal programme
record. For instance, lake deposits are known to be favourable for
for support of leading scientific schools, RFFI grant N 00-15-97934.
insect burial in contrast to deposits left by running waters. If a group of
Participation of M.B. Mostovski was helped by Dr. J. Ansorge
insect fossils has a poor record in spite of being aquatic ( judging from
(Institut für Geologische Wissenschaften, Greifswald), Dr. B.R.
its morphology: otherwise habits are unknown), it most likely occurred
Stuckenberg (Natal Museum, South Africa), Dr. E.A. Jarzembowski
in streams, and almost certainly so in cases where the few available
(Maidstone Museum and University of Reading, UK), Mr. R. Coram
fossils are worn and incomplete in contrast to other aquatic fossils col-
(University of Reading, UK), Dr. X. Martínes-Delclòs (Universitat de
lected there.
Barcelona, Spain), Dr. A. Arillo (Universidad Complutense, Madrid,
Palaeontology is an important, though not the only, way to trace the
Spain), and was supported in part by the International Science
history of insects: phylogenetics is another and is equally important,
Foundation, the Palaeontological Society (USA, grants nos. RG0-638
and it is noteworthy that it is largely independent of the fossil record in
(B) and RG0-822-7) and the Geologists’ Association (UK), ESF
its sources and inferences. Both ways are widely used and are mutually
Project “Fossil Insects Network”.
helpful and exert controls on one another. Using the words of Willi
Studies on terrestrial and aquatic insect palaeoecology
Hennig, they provide mutual illumination. There are even quantitative
(†V.V. Zherikhin and N.D. Sinitshenkova, respectively) have been
tools in progress aimed at helping to assess how well the fossil record
partially supported by the European Science Foundation (project
and various phylogenetic hypotheses agree with one another (e.g. the
“Fossil Insects”) and the Federal Scientific and Technical Programs
ghost range method, RASNITSYN 2000a).
(programmes “Evolution of the Biosphere” and “Co-evolution of
Phylogenetic research is popular and widespread, mainly in the
Ecosystems”).
form of cladistics (particularly since its fundamentals were rediscov-
Unless stated otherwise, illustrations are provided by the authors of
ered by the English speaking audience in HENNIG 1966a). However,
the respective chapters. M.K. Emelyanova (Palaeontological Institute
this is not the case for palaeoentomology, because palaeoentomologists
RAS, Moscow) helped in the formatting and the enhancing of many
were and are still few in number, and are widely scattered, except for
insect restorations and photographs. Thanks also go to Mrs. I.V.
the Moscow research group with its associates, which has been the
Kistchinskaya (Kishchinskaya) (Data+, Ltd.) for help in preparing the
most prolific and productive such group in the world since the 1960s.
maps of the fossil sites (see Figs. 3–5) with the use of the Arc View GIS
So it is not completely by chance that the present book is written
software programme.
mainly by those Russian scientists, but importantly it attempts to cover
all the world’s material and all the available information. Alexandr P. Rasnitsyn and Donald L.J. Quicke
xi
Contributors
Natalya V. Belayeva, Chair of Entomology, the Moscow State Alexandr G. Ponomarenko, Palaeontological Institute RAS, Moscow
University, Vorob’evyGory, Moscow, 119899 Russia, 117868 Russia, [email protected].
[email protected].
Yuri A. Popov, Palaeontological Institute RAS, Moscow 117868
Vladimir A Blagoderov, Palaeontological Institute RAS, Moscow Russia, [email protected].
117868 Russia, currently Department of Entomology, American
Ludmila N. Pritykina, retired from Palaeontological Institute RAS,
Museum of Natural History, Central Park West at 79th Street,
Moscow 117868 Russia, [email protected].
New York, NY 10024-5192, USA, [email protected].
Donald L.J. Quicke, Department of Biology, Imperial College at
Viktor Yu. Dmitriev, Palaeontological Institute RAS, Moscow 117868
Silwood Park, Ascot, Berkshire SL5 7PY UK, [email protected].
Russia, [email protected].
Alexandr P. Rasnitsyn, Palaeontological Institute RAS, Moscow
Kirill Yu. Eskov, Palaeontological Institute RAS, Moscow 117868
117868 Russia, [email protected].
Russia, [email protected].
Dmitry E. Shcherbakov, Palaeontological Institute RAS, Moscow
Andrey V. Gorokhov, Insect Taxonomy Laboratory, Zoological
117868 Russia, [email protected].
Institute RAS, University Embarkment 1, St Petersburg 199034
Nina D. Sinitshenkova, Palaeontological Institute RAS, Moscow
Russia, [email protected].
117868Russia, [email protected].
Vladimir D. Ivanov, St PetersburgState University, University
Sergey Yu. Storozhenko, Biological and Pedological Institute, Far
Embarkment 7, St Petersburg199034 Russia, [email protected].
East Branch RAS, Vladivostok 690022 Russia, entomol@online.
Nikita Ju. Kluge, St Petersburg State University, University
marine.su.
Embarkment 7, St Petersburg 199034 Russia, [email protected].
Irina D. Sukatsheva, Palaeontological Institute RAS, Moscow 117868
Mikhail V. Kozlov, Laboratory of Ecological Zoology, University of
Russia, [email protected].
Turku, Turku FIN 20500 Finland, [email protected].
Valentina N. Vishniakova, retired from Palaeontological Institute
Elena D. Lukashevich, Palaeontological Institute RAS, Moscow
RAS, Moscow 117868 Russia, [email protected].
117868 Russia, [email protected].
Peter Vršanský, Palaeontological Institute RAS, Moscow 117868
Mikhail B. Mostovski, Palaeontological Institute RAS, Moscow
Russia, [email protected].
117868 Russia, [email protected].
†Vladimir V. Zherikhin, Palaeontological Institute RAS, Moscow
Viktor G. Novokshonov, Perm State University, Bukireva 15, Perm
117868 Russia.
614600 Russia, [email protected].
xii
1
Introduction to Palaeoentomology
1.1
very useful tools, or that groups should not be clustered on the basis of
Scope and Approach shared derived characters (synapomorphies) wherever possible.
Rather, many believe that phylogenetic inferences are exactly the same
A.P. RASNITSYN as all other scientific statements in that they can never be proved nor
rejected, but are for ever destined to persist as more or less likely
hypotheses. Indeed, the inaccessibility of the finalverification of a sci-
entific statement has become almost a truism in post-Popperian times,
and yet the final falsification of one statement is nothing more than the
1.1.1. COVERAGE
verification of an alternative statement, that is, the falsifying result is
This book tries in general to cover the whole extent of the history of neither by chance nor due to unconsidered influences or circum-
insects in time and space in detail (Fig. 1). The exception is the stances. As a result there is no single best method for inferring phylo-
Quaternary Periodwhich is only occasionally considered here in depth genies, such as the outgroup approach comparison praised by some
because Quaternary palaeoentomology is a specialised field with its authors. Contrarily, many methods are good, each in its own place, and
own, rather specific methods and approaches. Quaternary palaeoento- their application can be efficiently regulated in the form of presump-
mology has recently been dealt with in considerable detail by ELIAS tion. Thus much of the work described in this book relies on making
(1994), and this might excuse our omission here. We understand that presumptions. This is not a trivial matter because of the often incom-
the selection of which subjects to present and discuss here reflects our plete nature of the evidence that scientists in general, and palaeontolo-
personal views and tastes, and that many important subjects may have gists in particular, have to deal with. A presumption is a statement
been missed, either by mistake or ignorance. We hope that our readers based on observations that a particular kind of result occurs more com-
will help us to make any future editions better than the present one. monly in particular circumstances, and thus it is to be considered as
The geochronological scale employed here is shown in Fig. 2, and most likely irrespective of the existence of confirmatory evidence, but
many insect fossil sites are displayed on the maps (Figs. 3-5). The not, of course, in the presence of reasonably sound contrary evidence
sites/site groups numbered and labelled there are the same as those (RASNITSYN 1988a, I992a, 1996, RASNITSYN & DLUSSKY 1988). For
catalogued in Appendix (Chapter 4). instance, we should consider any similarity between organisms as
The class Insecta is taken here in its narrow sense, that is, not includ- inherited from a common ancestor and not gained independently (as
ing the entognathous orders (Acerentomida = Protura, Campodeida = homoplasy), unless and until strong contrary evidence is presented.
Diplura, Podurida = Collembola). Although the relationships of these This has been termed the presumption of cognisability of evolution,
three orders is not fully resolved, there is growing evidence that they and it is equivalent to the auxiliary principle of HENNIG (1966a).
may form a monophyletic group together with the myriapods Another type of presumption that is particularly relevant to palaeontol-
(REMINGTON 1955, HANDSHIN 1958,MELNIKOV 1974a,b,RASNITSYN ogy (the palaeontological presumption) is that of two apparently
1976, MELNIKOV & RASNITSYN 1984, D. Shcherbakov, submitted) as closely related groups, the one entering the fossil record earlier should
explained in some detail below. be considered as ancestral unless and until sound contrary evidence is
presented. Likewise, we can recognise a biogenetic presumption, that
is, a transformation series should be polarised in agreement with the
ontogenetic succession of the respective character states; the outgroup
1.1.2. PHYLOGENETIC APPROACH
presumption, that a character state found only within a group should
It may come as a surprise to many members of the western scientific be considered apomorphic in respect to that found both within and out-
community that cladistics has not been universally accepted as the one side the group, and of course, the presumption of parsimony, i.e. that
and only method to be applied to classifying organisms by entomolo- the most likely cladogram is that one necessitating the least number of
gists and other systematists of the former Soviet Union. That is not to honnoplasies (i.e. the most parsimonious one). All of these presump-
say that cladistics and parsimony analysis are not regarded as being tions are accepted below unless there is convincing evidence to the
l