Mesozoic
Coleoptera

L V ARNOL'DI

V V ZHERIKHIN

L M NIKRITIN

A G PONOMARENKO

This monograph is a major treatment
of Mesozoic Coleoptera, being the
first review of the evolution of beetles
during that era. The Early Mesozoic,
Late Mesozoic and Late Cretaceous
stages are recognized in the phylo-
genesis of Mesozoic Coleoptera.
Special attention has been paid to
their ecological associations, parti-
cularly with food plants. One hundred
and eleven species, 64 genera and 19
families are described, of which all
species, most of the genera and four
families are new.

This work is of interest to ento-
mologists and paleontologists,
especially paleobotanists and strati-
graphists of Mesozoic continental
deposits.

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Mesozoic Coleoptera

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Mesozoic Coleoptera

L.V. Armol’di, V.V. Zherikhin,
L.M. Nikritin and A.G. Ponomarenko

Scientific Editor
Natalia J. Vandenberg

Smithsonian Institution Libraries
and
The National Science Foundation
Washington, D.C.
1992

TT 81-52002

Mezozoiskie Zhestkokrylye. Akademiya Nauk SSSR,
Trudy Paleontologicheskogo Instituta, Vol. 161.

Nauka Publishers
Moscow, 1977

Translated from the Russian
General Editor: Dr. V.S. Kothekar

© 1991 Oxonian Press Pvt. Ltd., New Delhi

Library of Congress Cataloging-in-Publication Data
Mesozoic Coleoptera.

Translation of: Mezozoiskie zhestkokrylye.

Bibliography: p.

1. Beetles, Fossil. 2. Paleontology—Mesozoic.
I. Arnol’di, L. V. (Lev Vladimirovich) II. Vandenberg,
Natalia J. III. Title.
QE832.C6M4913 1989 565’.76 88-607938

Translated for the Smithsonian Institution Libraries, pursuant to an
agreement with the National Science Foundation, Washington, D.C., by
Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001

Typeset by Cristina Graphs Pvt. Ltd., Delhi and printed at
Pauls Press, New Delhi, India

This monograph is a major treatment of Mesozoic Coleoptera, being the first
review of the evolution of beetles during that era. The Early Mesozoic, Late
Mesozoic and Late Cretaceous stages are recognized in the phylogenesis of
Mesozoic Coleoptera. Special attention has been paid to their ecological as-
sociations, particularly with food plants. One hundred and eleven species,
64 genera and 19 families are described, of which all species, most of the
genera and four families are new.

This work is of interest to entomologists and paleontologists, especially
paleobotanists and stratigraphists of Mesozoic continental deposits.

Plates 14, illustrations 109, bibliography 58 entries.

Editor-in-chief
B.B. Rohdendorf

Doctor of Biological Sciences

hil
pbs

Foreword to the English Edition

The Smithsonian Institution Libraries, in cooperation with the National Sci-
ence Foundation, has sponsored the translation into English of this and
hundreds of other scientific and scholarly studies since 1960. The program,
funded with Special Foreign Currency under the provisions of Public Law
480, represents an investment in the dissemination of knowledge to which
the Smithsonian Institution is dedicated.

The present edition is a translation of a major collaborative work on
Mesozoic fossil beetles written by four eminent Soviet scientists. This is the
first time that detailed descriptions based on the exceptionally fine collec-
tion of the Paleontological Institute, Academy of Sciences, USSR, have
been made accessible to English-language readers. The Institute collection
contains over 15,000 specimens of fossil beetles from the Triassic, Jurassic
and Cretaceous, including many key examples exhibiting a remarkable state
of preservation.

The Mesozoic Era marks a crucial period in the evolution of beetles: at
its commencement we encounter some of the first examples of specimens
possessing characteristics suggestive of particular suborders of Recent
Coleoptera, and by its close, many modern families may be found among its
diversified fauna. The scope and importance of the present work is discussed
by B.B. Rohdendorf in the brief abstract and foreword to the Russian
edition, and will not be repeated here. However a few comments relevant
to the English-language edition are in order.

The translated version of Mesozoic Coleoptera has remained as faithful
as possible to the original work, but what we have attempted here is a
translation of ideas, not of words. In many instances a departure from a more
verbatim style was necessary in order to preserve meaning. This was
particularly true of the narrative portions of this work where we took the
liberty of breaking up some of the longer sentences and, occasionally, of
rearranging the sequence of ideas. Sometimes additional information which
had been implied but not strictly stated in the Russian text was added to the
translated version either in brackets or as a footnote. Russian terms which
represent a translation of Greek or Latin taxonomic categories were treated
as equivalent to the latter. For example, the term “zhestokokrylye” was

Vili

translated as the scientific ordinal name “Coleoptera,” not as “sheath-
wings” (a literal translation) or “beetle” (the common name). Proportional
measurements are reported in a format which we hope will make them more
comprehensible to English speakers; their treatment is discussed in a brief
footnote at the beginning of the Descriptions of taxa section.

The English-language edition of Mesozoic Coleoptera was initiated at
the request of Terry L. Erwin, Curator of Coleoptera at the National
Museum of Natural History, Washington, D.C., and published by the
Smithsonian Institution Libraries in cooperation with the National Science
Foundation. Many people have contributed towards the preparation of this
publication. In addition to the key individuals whose names appear within
the covers of this book, I would like to thank the many people who have
helped with the translation, coordination and production of this edition. I
would also like to thank my father, George N. Vandenberg, for his assistance
in interpreting some of the Russian idioms, and my colleague Alan I. Kaplan
for the loan of pertinent literature from his private library.

July 1989 Natalia J. Vandenberg, Ph.D.
Scientific Editor
Albany, California

3

nS

Foreword

Coleoptera are distinguished from other organisms by their exceptional
diversity. The number of present-day species in this order not only exceeds
that of any other order, but of all plants and animals combined, excluding
insects. Beetles play a significant ecological role and are economically
important as crop and stored-products pests. Our present knowledge of the
structure, ecology and taxonomy of Coleoptera is fairly advanced, but
information on their evolution (i.e. actual stages in the formation of the
group) is highly inadequate. Study of the geological history of beetles is
important not only for a description of their phylogeny and classification,
but also, in a broader sense, for understanding many trends in the evolution-
ary process.

From the end of the Paleozoic era, Coleoptera have played a vital role
in continental biocenoses. Moreover, in the Early Mesozoic their role was
probably even more important than at present. Data on ancient beetles are
also important for an understanding of trends in the formation and function-
ing of ancient biocenotic complexes. At present, our knowledge of ancient
beetles is totally inadequate. Almost all information pertains to Cenozoic
beetles which are poorly distinguished taxonomically from present-day
forms, belonging to the same families and, in many instances, even to the
same genera. The entire protracted period of their early history—the end of
the Paleozoic and particularly the whole Mesozoic—remains inadequately
studied to date. This has generated increased interest in ancient Coleoptera,
particularly Mesozoic ones.

Entomological studies conducted since the 1920’s on a variety of
Mesozoic faunal complexes in the USSR have provided abundant coleop-
teran material. This material has been greatly enriched in the last decades
and, more importantly, its quality has vastly improved since attention
focussed on the search for intact whole fossils during field studies. Most
early descriptions of Mesozoic Coleoptera were based on the study of a
single elytron or other fragments; these were not sufficient to permit an
exact determination of the insects’ taxonomic affinities. Present-day studies
of complete fossils of ancient Coleoptera have made it possible to reliably
recognize their characteristics. Following the study of individual groups of
beetles, conducted by specialists of each order, a detailed account of

x

Mesozoic beetles could be compiled together with outlines of the faunal
complexes of Mesozoic Coleoptera.

The review of Mesozoic Adephaga is particularly important and in-
cludes descriptions of 56 species, 31 genera and 8 families. Literally for the
first time, phylogenetic trends within the suborder are evaluated using
primarily new material, and taking into account diverse ecological and
functional characteristics (A.G. Ponomarenko). This matter is considered
relatively fully in the description of Adephaga. The large suborder Polyph-
aga, on the other hand, is treated more unevenly. For example, the unexpect-
edly large chapter on the Rhynchophora (33 species, 18 genera, 3 families),
specialized beetles which are usually placed at the end of the system even
though they are rather well represented in the Mesozoic material (L.V.
Arnol’di and V.V. Zherikhin). Important are the descriptions of some Early
Cretaceous Scarabaeidae (L.M. Nikritin) which until recently were practi-
cally unknown from the Mesozoic. Finally, some phylogenetically interest-
ing Polyphaga are described, for example, the aquatic Hydrophiloidea (8
species, 5 genera, 2 families); the taxonomic positions of many of these are
uncertain (A.G. Ponomarenko). Equally interesting are taxonomically |
isolated Cucujiformia (3 families) that permit some generalization (V.V.
Zherikhin). On the whole, this volume is a major new treatment of a widely
distributed and important group of Mesozoic insects. It allows the develop-
ment of various conclusions and comparisons regarding ecology and
evolution of the continental fauna of the earth.

B.B. Rohdendorf

Contents

Russian English

page™ page
FOREWORD TO THE ENGLISH EDITION o Vil
FOREWORD Sar ix
INTRODUCTION—A.G. Ponomarenko SSxidags) 1
COMPOSITION AND ECOLOGICAL CHARACTER-
ISTICS OF MESOZOIC COLEOPTERA—
A.G. Ponomarenko at eS 5
DESCRIPTIONS OF NEW TAXA sealla l/ 19
Suborder Adephaga—A.G. Ponomarenko eel ay 19
Family TRIAPLIDAE Ponomarenko,
fam. nov. spall 19
Infraorder Hydradephaga ss 23
Family PARAHYGROBIIDAE Ponomarenko,
fam. nov. Be) We) 78
Family COPTOCLAVIDAE Ponomarenko,
1961 ee oP 26
Family LIADYTIDAE Ponomarenko,
fam. nov. aS 48
Family DYTISCIDAE Leach, 1815 .. 40 52
Family GYRINIDAE Latreille, 1840 . 42 54
Infraorder Geadephaga .. 46 60
Family TRACHYPACHEIDAE Leconte,
1861 . 46 60

*The page numbers of the Russian original appear in the left-hand margin of the text—
General Editor.

xii
Russian English

page page
Family CARABIDAE Latreille, 1802 cresyl 96
Adephaga Incertae Sedis 90 122
Mesozoic Stages in the Evolution of
Adephaga—A.G. Ponomarenko 96 130
Suborder Polyphaga .. 104 142
Polyphaga Incertae Sedis—A.G. Ponomarenko .. 105 143
Infraorder Staphyliniformia—A.G. Ponomarenko .. 106 145
Family HY DRAENIDAE Mulsant, 1844 : 106 145
Family HY DROPHILIDAE Leach, 1815 .. 108 148
Family SILPHIDAE Latreille, 1807 Solely 159
Infraorder Scarabaeiformia—L.M. Nikritin sg bD 162
Family SCARABAEIDAE Latreille, 1802 su UL 162
Infraorder Elateriformia—V.V. Zherikhin .. 130 177
Family CEROPHYTIDAE Latreille, 1834 .. 130 177
Series of Superfamilies of Cucujiformia—V.V.
Lherikhin SS) 182
Family ACANTHOCNEMIDAE Crowson,
1964 se NSIS) 182

Family CRYPTOPHAGIDAE Erichson, 1845 .. 138 187
Family LATHRIDIIDAE Redtenbacher, 1845 .. 140 189

Rhynchophora—L.V. Arnol’ di .. 142 191
Family EOBELIDAE L. Arnol’di, fam.
nov.—L.V. Arnol’ di . 144 195
Family ATTELABIDAE Billberg,
1820—V.V. Zherikhin ye WTA) 242
Family CURCULIONIDAE Latreille,
1802—V.V. Zherikhin A178 244
BIBLIOGRAPHY 83 251

PLATES a WEY PE

Introduction

5 Ofall stages of the historical development of beetles the Mesozoic is the most
difficult to study. Paleozoic beetles, whose remains are found only in the
Permian deposits, were apparently rather less numerous and diverse. So far
only 200 or so coleopterous remains have been collected from the Permian
continental strata. The remains of the Permian beetles are much more
abundant in regions thought to have had a temperate climate. No Permian
beetles have been found, which could be assigned with certainty to the more
advanced suborders. All of them either definitely are members of the sub-
order Archostemata or can belong there. To date, 32 genera of Permian
Coleoptera belonging to 5S families have been described. It is true that most
of them have been established from a single elytron, however the evolution-
ary characteristics of Permian beetles enable us to consider the taxa estab-
lished from single elytra as fairly natural (for further details see Pono-
marenko, 1969). The limited diversity of Permian beetles and the fact that
their evolution can be followed even from isolated sclerites, simplifies the
task of describing them and of studying their historical development.

The study of Cenozoic beetles has been similarly facilitated although for
another reason. Cenozoic Coleoptera differ very little from extant forms,
belonging to the same families and, in many instances, even to the same
genera. This makes it possible to differentiate them using indirect diagnostic
features which are often more highly visible on fossil remains than are the
key characters that accurately establish their taxonomic position.

The study of Mesozoic beetles is a more complex problem. On the one
hand, the Mesozoic beetles are almost as diverse and numerous as the extant
beetles. At any point of time in the Late Mesozoic, these beetles were
certainly represented by hundreds of thousands of species belonging to over
a hundred families (there are almost 200 families of extant beetles). Thus in
a single Jurassic site of Karatau, the family Cupedidae is represented by 20
species (Ponomarenko, 1968b), Eobelidae by 30 species (Arnol’di, this
monograph), and Elateridae by more than 20 genera (Dolin, 1975) with
perhaps about 100 species.

On the other hand, most Mesozoic beetles belong to extant families or
those closely related to them. However, representatives of the contemporary
families usually differ so markedly from Cenozoic* ones that the use of

*From the context it seems likely the word “Mesozoic” was intended—Scientific Editor.

a

2

indirect diagnostic features becomes unreliable. Intact and well-preserved
remains are therefore a prerequisite for determining the taxonomic position
of Mesozoic beetles since important taxonomic characters can be studied in
them (e.g., the structure of antennae, propleura, base of the abdomen, hind
coxae, and the number and arrangement of abdominal spiracles).

Unfortunately, practically none of the earlier described remains of
Mesozoic Coleoptera meet these requirements. For most of them, the
taxonomic position indicated by the author was not substantiated at all or was
substantiated only by indirect characters and external similarities. The
remains of an excellently preserved beetle, Proteroscarabeus yeni Grabau,
were studied by two researchers (Grabau, 1923; Ping, 1928). All the same,
the description contains no characters that formally justify its inclusion in the
family Scarabaeidae. Hence the Mesozoic history of beetles remains almost
totally unexplored even though nearly five hundred beetles have been
described from this era.

The foregoing explains the interest in almost any publication dealing
with fairly complete remains of Mesozoic beetles. Particularly important is
the study of the vast collection of Mesozoic beetles preserved in the Paleon-
tological Institute, Academy of Sciences, USSR. This collection includes
over 15,000 remains of Triassic, Jurassic and Cretaceous beetles and evident-
ly exceeds all other collections of Mesozoic beetles combined. This collec-
tion includes many examples of exquisitely preserved ancient beetles in
which not only integumental structures, but also other characters rarely
available to the paleontologist can be studied (e.g., the structure of the hind
wings, mouth parts and endoskeleton). These latter characters permit
speculation about specific musculature.

Unfortunately, this magnificent collection remains poorly studied. Ear-
lier, only archostemate beetles have been investigated in great detail
(Ponomarenko, 1964, 1966, 1968a, b, 1969) and three other papers have been
devoted to major groups of beetles from the Jurassic site of Karatau
(Tikhomirova, 1968; Medvedev, 1968; Dolin, 1975). The remaining papers
contain descriptions of only individual forms of Mesozoic beetles.

The present work, as well, does not present an exhaustive description of
Mesozoic Coleoptera, even of those solely from the collections of the
Paleontological Institute. Only Adephaga were examined in considerable
detail. From this suborder, 58 species covering 32 genera and 8 families are
described below. Three families, 28 genera and all the species are new. Aside
from the Mesozoic beetles described here, less than 10 genera are known
which can be definitely included in the Adephaga. Most of the taxa in the
present book are described from adult beetles and only two species from
larvae. However, the volume of material on these two larvae is almost as
large as that on the adult beetles: in all, nearly 150 adults and 100 larvae.
Based on the Mesozoic adephagan material studied, it was possible to

3

describe in general terms the characteristics of the Mesozoic stage in the
evolution of these beetles.

In this work, beetles of the suborder Polyphaga are presented in in-
dividual groups. Only a small proportion of the representatives from this
gigantic group, present at the Paleontological Institute, have been described.
However, some tentative ideas can be developed about the Mesozoic evolu-
tion of Polyphaga using the newly described material combined with earlier
descriptions. Only two series of superfamilies recognized by Crowson (1971)
are not represented in the Mesozoic: Eucinetiformia and Bostrychiformia.
The former is undoubtedly present in the Jurassic of Karatau and is repre-
sented by forms very closely related to the extant ones. The latter obviously
existed in the Mesozoic; its remains must be available in collections, but,
have not yet been recognized. In all, 53 species of Polyphaga belonging to
32 genera and 11 families are described below.

Some of the beetles are described from inclusions in Mesozoic fossil
resins, often called ambers by analogy with Baltic amber. Beetles embedded
in Mesozoic resins have been reported earlier (Carpenter, et al., 1937;
McAlpine and Martin, 1969) but remain undescribed. A study of the remains
of ancient beetles in the transparent resins permits their description with the
same degree of detail as for the extant forms. It is a very important observa-
tion that within the resins, as arule, one encounters minute beetles which are
difficult to study from their impressions. Resins, thus considerably expand
the range of information available to us on ancient insects.

Different groups of Mesozoic beetles are described here by four authors
who have attempted, insofar as possible, to achieve a uniform style of
presentation. However, existing traditions in the style of descriptions, and
even in the morphological nomenclature of different groups, preclude com-
pleie uniformity. All the described material is preserved in the Paleontologi-
cal Institute, Academy of Sciences, USSR (PIN). Some important principles
in the study of fossil beetles have been stated earlier (Ponomarenko, 1969b)
and do not warrant rediscussion. The exact taxonomic positions are far from
being established for many of the described forms. Such forms have been
placed in a separate genus only when substantial evidence indicated that they
could not beiong to any extant or extinct genus. When such a demonstration
was not possible, the remains were described as a formal collective genus
named after the type genus of the family with the suffix “ites”. In some
instances, formal genera were proposed for fragmentary fossils. Since a
formal comparison of these forms is not possible, the section “Comparison”
in their description has been replaced by “Taxonomic position” to discuss
the probable taxonomic status of the fragmentary form. The material studied
originates from sites repeatedly described and discussed in the literature.
Hence they have not been described here; their description can be found in
earlier papers (Ponomarenko, 1969; Rasnitsyn, 1969, 1975).

The main faunal complexes of Mesozoic coleopterans are characterized
in a very brief and concise form. Unfortunately, their analysis was possible
only for the territory of the USSR. The rather numerous fossil beetles known
from other territories are almost totally inadequate for comparison because
of imperfect descriptions. Naturally the documented Mesozoic history of

beetles is fragmentary and does not give acomplete picture of the Mesozoic.

evolution of beetles; it presents only individual moments of this history.

Composition and Ecological Charac-
teristics of Mesozoic Coleoptera

8 As previously mentioned, the sites of Mesozoic beetles are very unevenly
distributed over the earth. Rich sites, yielding an abundance of suitable study
material, are even more unevenly distributed. A large proportion of the fossils
of ancient Coleoptera consist of isolated elytra; but, unfortunately, in the
Mesozoic these only rarely provide a basis for ascertaining the taxonomic
position of the beetle. In the future, when our knowledge of ancient beetles
has increased considerably, it will be possible to determine the taxonomic
position of incomplete fossils and use them for paleobiogeographic and
biostratigraphic purposes. For the present, well-preserved fossils are ab-
solutely necessary. Only 1-10% of the fossils collected from various sites
are useful for study, but hundreds and thousands of specimens from a given
site are necessary for any valid judgement on beetle complexes.

‘ The potential that any site offers for analysis depends greatly on its
taphonomy. In some sites, more or less intact beetles are preferentially buried
and sometimes the number of entire impressions may even exceed the
number of isolated elytra. In other sites the remains get dislodged and
dismembered before burial so that the number of entire specimens is reduced
to only a small percentage. The first type of site is highly promising for
investigation, but when dealing with the second type of site, thousands of
remains need to be collected in order to obtain the required material. Even
collections of many isolated elytra are not entirely useless. Each separate
elytron may not be identifiable but the overall spectrum of morphological
types of elytra and their quantitative relations often turn out to be highly
significant.

Few of the Mesozoic sites have a potential value of the study of ancient
Coleoptera. This is despite the fact that during the Mesozoic, beetle remains
are most numerous in oryctocoenoses, and the Mesozoic fossil insect sites
number in the hundreds. The problem is well illustrated by a recent attempt
to re-examine the type material from the well-known site of Mesozoic insects
at Solenhofen in the Federal Republic of Germany (Ponomarenko, 1971a).
The preservation of the fossil remains in this site has always been considered
exemplary but in the studied collection, including a large proportion of type
material, there were hardly any remains whose taxonomic position could be

\o

6

decided with absolute conviction. In most cases it had to be simply stated
that due to the poor state of preservation they were not suitable for the present
investigation. The taxonomic assignments made by the original authors
proved totally incorrect. Only one species has been found to actually belong
to the family in which it was placed in the original description.

Presently almost all the sites yielding abundant specimens of ancient
beetles are located in Asia, mainly in the northern part. There are very few
rich sites on other continents. The abundance of material collected in the
Asiatic part of the USSR is due to two reasons. First, only in the USSR have
expeditions recently been conducted by a large team of paleoentomologists
for the sole purpose of collecting fossil insects. The collections from the
USSR surpass the remaining global collections. Second, the inequality of
sites in the European and Asiatic parts of the USSR arises from differences
in their geologic structure. The nature of deposits in these territories is such
that Mesozoic fossil insects are very rare in the European part of the USSR,
whereas the Paleozoic collections from both territories are of the same order.
Western Europe is far more favorable for the Mesozoic insect fossils but
practically no special expeditions have been organized solely for fossil
collection.

The faunal complexes discussed here appear rather geographically
restricted and do not represent the entire global biota of the Mesozoic era. At
the same time, the sites examined occur in different climatic or, in any case,
floristic zones. The chronologically sequential sites are frequently found in
different zones and thus it is difficult to determine the cause of such
differences—their position in different zones, or evolutionary shifts. These
factors inevitably influenced the nature of the investigation of the fauna
described below.

The oldest Mesozoic faunal complexes—the Triassic and early Jurassic
ones—sharply differ from the later complexes. In composition they are closer
to the Paleozoic than to the Late Mesozoic. Almost all their constituent
beetles belong to the most ancient and the most primitive suborder Archo-
stemata. Unfortunately, beetles from the Early Triassic are extremely rare.
The rich Dzhailyaucho site in southern Fergana was considered Early Trias-
sic, and paleobotanists (Sikstel’, 1962, 1965) even assigned its lower strata
to the Permian. Recently I.A. Dobruskina (1970, 1974) showed that all the
deposits in the Madygen area, including even Dzhailyaucho, are coeval and
belong to the end of the Middle—Early Upper Triassic. Archostemata have
already been described from Dzhailyaucho; astonishing differences have
been observed between these supposedly Early Triassic beetles and the Late
Permian ones, in addition to their similarity with Coleoptera from the known
Middle and Upper Triassic sites (Ponomarenko, 1969b). A similar type of
resemblance was reported by A.P. Rasnitsyn (1969) for Hymenoptera. This
was ascribed to the fact that the Triassic fauna of Australia, where rich

10

7

collections of Late Triassic insects are known, has amore ancient continental
character than that from the northern continents. The need for such supposi-
tions no longer exists.

As mentioned earlier, almost all Triassic beetles are Archostemata. More
than 15,000 insect fossils have been collected to date from Triassic deposits
in the Madygen area. Beetles occupy third place after cockroaches and
cicadas. Nearly 3,500 fossil beetles have been recovered. It is difficult to
establish the exact proportions of beetles of different suborders. Isolated
elytra constitute the vast majority of the fossil remains of beetles from this
site. Using these remains, it is not possible to distinguish the schizophoroid
Archostemata from primitive Adephaga, or Ademosynidae (related to Ar-
chostemata)* from many Polyphaga. A comparison of groups must therefore
be made from only a few well preserved beetles. To date, a total of 63 species
(here and throughout including those described in the text) of Triassic beetles
of 34 genera and 7 families have been described from Madygen. Of these,
only four species definitely do not belong to Archostemata. True, some
beetles belonging to Adephaga were possibly described as Archostemata.
The difficulties encountered in differentiating the Mesozoic members of
these beetles are discussed in the section specially devoted to the Mesozoic
Adephaga, hence they are not discussed here. However, in the Triassic faunal
complexes, both higher suborders**, even with the most liberal estimate, do
not make up the 10% estimated earlier (Ponomarenko, 1968a).

Among the Triassic Archostemata, members of the family Cupedidae
are the most varied. They constitute 25% to 50% of all beetles at the various
sites. From the Dzhailyaucho site alone 29 species have been described, but
their diversity is by no means fully revealed. There are numerous beetle
fossils that clearly do not belong to these species and remain unidentified due
to their poor preservation, which precludes their formal inclusion in any
particular genus. The Cupedidae first appeared in the Triassic (their fossils
are not known from older deposits) and are represented by all of the three
subfamilies, one of which is known only from the Triassic. In no other period
have the Cupedidae been so numerous and diverse; the cupedid taxa at
Dzhailyaucho far outnumber the extant taxa the world over.

The Schizophoridae with 16 species and 10 genera occupy the second
position. Although their species are far less than those of cupedids, they are
close to cupedids in the number of genera, and surpass them in the number
of extant forms. In morphological diversity, they are on a par with the
present-day ground beetles excluding the paussids. The remaining three
families—Tricoleidae, Ademosynidae and Catinidae—are represented by
relatively few and less diverse forms.

*The authors’ treatment of Ademosynidae is inconsistent. It is altemately placed within
Archostemata or contrasted with the latter—Scientific Editor.
**Presumably Adephaga and Polyphaga—Scientific Editor.

1

—

The suborder Adephaga is represented by the unique family Triaplidae,
possibly the most primitive member of the suborder. The presence of one
representative of Trachypacheidae, poorly differentiated from the Early
Jurassic forms, is unexpected. Hydradephaga are not found in Madygen, but
in the Upper Triassic sites of Garazhovka in the Ukraine an incomplete beetle
fossil has been found, which may be the form directly related to the whirligig
ancestors.

Representatives of the suborder Polyphaga are even less numerous. A
single form of uncertain affinities is described below which may be very
close to the Dascilloidea (sensu Crowson, 1971) and is characterized by many
primitive features. In addition there are several interesting yet undescribed
forms. One fossil beetle, based on the shape of its pronotum, almost certainly
belongs to Elateroidea. Unfortunately, the preservation is so poor that it is
not possible to determine its family, particularly whether it is Praelateridae
or Elateridae. Finally, three fossil beetles have been found in Madygen,
whose presence in the Triassic was very unexpected: these beetles are very
close to the Jurassic Eobelidae described in the text. They have a typical
Rhynchophora morphology with a rather long rostrum and an oxycorynoid
body with a distinct longitudinal ridge on the propleuron.

The ecological spectrum of beetles found in Madygen is also interesting.
Phytophages are primarily represented by the very abundant cupedids. The
Triassic cupedids differ little from the Jurassic ones; the most specialized
forms, as far as one can judge from the shape of the body, are most closely
related to them. Among the Jurassic cupedids there are even extant genera,
hence there is no doubt that the Jurassic and Triassic cupedids hardly differed
in mode of life from their present-day relatives. It would follow that they
lived in wood broken down by fungi, feeding on the fungi and the decaying
wood. All the Polyphaga of Madygen, except the Eobelidae, also thrived on
wood, but of a far more decomposed nature. The Eobelidae were probably
spermophagous. The parallel flourishing of the Eobelidae with cycadophytes
suggests that the larvae of these ancient weevil-like beetles lived in
cycadophyte fruits.

A large number of beetles from Madygen were in some way linked to
water. All possible arguments in support of an aquatic or riparian life for most
schizophoroid Archostemata have already been advanced (Ponomarenko,
1969). No new evidence has been presented either to support or contradict
this point of view. These forms were probably poorly specialized aquatic or
riparian predators and detritovores which inhabited both sides of the
coastline. Forms adapted to swimming are not found among them. There are
also typical terrestrial forms, differing little in external appearance from
ground beetles. They differed from the primitive ground beetles of the genus
Sogdodromeus (Trachypacheidae), found alongside them, in having distinct-
ly tuberculate sculpturing of the integument typical of the Archostemata. It

9

is possible that unidentified primitive ground beetles are presentamong them.
The feeding habits of the Triaplidae remain unknown. Their closest relatives,
the Haliplidae, are strict algivores, feeding on green and charophyceous
algae. The Triaplidae were probably also phytophages or detritivores as
indicated by the orthognathous or even opisthognathous head, whereas in
predators the head is almost always prognathous.

The largest fossil beds of Early Jurassic Coleoptera are also distributed
in Soviet Central Asia.* Unfortunately isolated sclerites, mainly elytra,
predominate in all the beds, and complete fossils are very rare. Although
several thousand beetle fossils have been collected, not many are suitable for
study.

As before, Archostemata predominate and presently constitute one-half
to three-fourths of all the beetles. Archostemata dominate also among the
described species: 22 species of Archostemata have been described versus
only 8 of the other beetles.

The Cupedidae, as usual, are the most widely distributed group of
Archostemata. However, they now constitute not more than 10% of all the
beetles, and only in one site, Kyzyl-Kiya, 26%. Triadocupedinae, the most
primitive subfamily of cupedids, has not been found in the Jurassic; it is true
that even the most advanced, primarily Cenozoic subfamily Cupedinae,
which existed in the Triassic, is also not found in the Early Jurassic. All the
cupedids known from the Early Jurassic belong to Ommatinae; the Mesozoic
tribe Notocupedini accounts for 75% of all the forms found. Only 7 species
of the genus Notocupes have been described from the Early Jurassic, but
elytral diversity suggests a much higher number. The extant cupedids, Omma
and Tetraphalerus, appeared in the Early Jurassic.

The remaining Archostemata are almost exclusively represented by
aquatic forms. Outwardly they strongly resemble water scavenger beetles
(Ademosynidae) and the rather diverse Schizophoridae. The latter retain
almost all the lineages present in the Triassic but their diversity is generally
markedly reduced. The number of terrestrial forms is particularly small. This
is probably explained by the displacement of such forms by the very similar
primitive ground beetles. Among aquatic schizophorids the appearance of
very characteristic Mesozoic forms with long, hooked legs and hypognathous
head is worthy of note. These beetles probably lived on the aquatic vegeta-
tion. One genus of the family Tricoleidae has also been reported.

*The Russian literature frequently refers to two distinct geographical localities “Tsentral’-
naya Aziya” (Central Asia) and “Srednaya Aziya” (loosely translated as Middle Asia). The
former term refers to Sinkiang Province in China (or the former Chinese Turkestan), while the
latter refers to the former Russian Turkestan (the Kirgiz, Uzbek, Turkmen and Tadzhik SSR).
Since there is no geographic locality by name Middle Asia, we prefer to call it Soviet Central
Asia—General Editor.

10

At the beginning of the Jurassic, beetles of the suborder Adephaga were
apparently represented by all the major Mesozoic groups, although all of
them are still not known. Whirligigs and primitive dytiscoids obviously must
have existed among the aquatic Adephaga, but in fact only one form,
Necronectulus, distinctly close to the dytiscoid root, is known. A primitive
fossil beetle of the family Coptoclavidae has been found in Soviet Central
Asia in the supposedly Early Jurassic deposits, but recently doubts have been
expressed about the correct age of these deposits: possibly they are younger.
In these deposits, terrestrial Adephaga are represented by fossils of Carabidae
and Trachypacheidae. These fossils represent the most primitive forms of
both families.

Well preserved fossil Polyphaga are rare in the Lower Jurassic sites. At
present only representatives of the Elateridae are known; or, more precisely,
its Mesozoic subfamily Photagrypninae and the family Praelateridae which
is closely related to click beetles (Dolin, 1973, 1975).

Unfortunately, the Middle Jurassic beetle fauna, next in chronology, has
been collected almost exclusively from sites in Siberia. Hence it is difficult

12 to say ifits lack of congruence with the preceding fauna is due to evolutionary
changes or simply to zoogeographic peculiarities. Moreover, the beetles in
these sites are represented only by aquatic or littoral* forms. Archostemata
are generally not found here. This is particularly significant with respect to
cupedids. Whereas other Archostemata may elude identification based on
isolated elytra, this possibility is ruled out for cupedids. In all the Siberian
Jurassic sites not a single cupedid elytron has yet been found. Hence their
abundance must be at least two orders of magnitude lower than in the Jurassic
sites of Europe and Soviet Central Asia. In Siberia, cupedid fossils were
discovered in the younger Cretaceous deposits. This difference is apparently
due to the characteristics of the Jurassic vegetation of Siberia, where gingkos
and chekans** predominated and conifers were rare.

Adephaga turn out to be the dominant group of beetles from these sites.
Even though the number of fossils collected here is not great, all the Mesozoic
families of aquatic Adephaga—Parahygrobiidae, Liadytidae, Coptoclavidae
and Gyrinidae—have been found in the Siberian sites whose ages are close
to Middle Jurassic. Fossils of the most primitive beetle larvae—larvae of
aquatic Parahygrobia, Angaragabus and Stygeonectes—have also been
collected from these deposits.They all belong to the dytiscoid Adephaga and
are hardly different from the extant dytiscoid larvae. Among them are
nektonic and benthic forms. These were all air breathers, with their well
developed tracheae modified to form a hydrostatic apparatus. One of the

*So given in the Russian original but by intent should read “aquatic, more particularly
littoral” —General Editor.
** Apparently representatives of cycadales—General Editor.

11

larvae (Angaragabus) showed a falciform mandible, lacking a retinaculum,
but with an internal canal. A larva of the staphyliniform Polyphaga, probably
a primitive hydrophilid, has also been found here.

It is noteworthy that to date no representative of Geadephaga has been
found at these sites. This may be considered a chance occurrence due to the
poor collections, but here, as well as in two other sites, fossils have been
found of a possible ecological analog of the Mesozoic ground beetles: the
primitive carrion beetle Mesagyrtes, closest to the extant predaceous carrion
beetles feeding on coastal mollusks. Fossils of this beetle constitute a third
of all the collected coleopteran fossils: an absolutely unique case. Only
primitive hydrophilids and staphylinids could be identified among the
remaining Polyphaga. This fauna is thus almost exclusively associated with
water, which makes its comparison with other fauna difficult. Interestingly,
this is notan isolated discovery; similar fauna have been collected from more
than ten sites spread over a vast territory from west Siberia to Trans-Baikal.

Beetles from the very end of the Jurassic are best known from the famous
Karatau site in south Kazakhstan. The site is a chain of rock outcrops
apparently formed in a large lake. The composition of insect fossils varies
somewhat in different outcrops, but whether this is due to variations in the
conditions of burial or the age of the deposits is not yet clear. More than
18,000 insect fossils have been collected from this site, also a few fossils of
other arthropods and a rather large number of vertebrates, particularly fish.
The collection from Karatau has more than 5000 fossil beetles. Since in
recent years only the intact, well preserved fossils have been collected, in
reality the number of beetle fossils could be very large*.

The composition** of beetles from Karatau differs sharply from that at
the beginning of the Jurassic. The Archostemata constitute only about 10%
of the beetles. Precise estimates give a high figure but this is probably an
overestimate since the elytra of cupedids were collected fully, but not those
of other beetles owing to poor preservation. Thirty species of Archostemata
representing 13 genera and 4 families have been described from Karatau.
Recent collections from this area almost doubled the number of beetles but
mainly repeated previously documented finds, yielding only two or three new
species. According to a preliminary estimate, the Archostemata constitute
approximately 5 to 10% of all taxa; and 75% of the Archostemata are
cupedids, which are particularly diverse in this site. An entire series of
endemic genera were restricted to the Late Jurassic or appeared only in the
Early Cretaceous; these existed alongside the genera which had appeared
earlier and persisted throughout the Cretaceous, a few even continuing to the
present day. The remaining Archostemata—from the families Ademosyni-
dae, Schizophoridae and Catiniidae—are unique forms and, judging from

*Not a verbatim but meaningful translation—General Editor.
**Species composition—General Editor.

12

their body structure, are aquatic. Among them, representatives of the genus
Tersus distinctly predominate. Judging from the long hooked legs, they
inhabited aquatic vegetation. The large number of specimens collected may
provide additional evidence of their aquatic life.

The number of accurately determined specimens of Adephaga exceeds
that of Archostemata, though their actual numbers were apparently similar.
Twenty-three species, 13 genera and 4 families of Adephaga have been
described from Karatau. The Hydradephaga are represented by one genus of
whirligigs and four genera of coptoclavids. The coptoclavid diversity is
greater here than in any other site. The Geadephaga are almost twice as
numerous as the Hydradephaga. The Carabidae and Trachypacheidae are
almost identically represented both in the number of taxa described and the
number of specimens.

Almost all Karatau beetles belong to Polyphaga. This is the most
primitive fauna in which the Polyphaga are distinctly dominant and very
diverse. Apparently, the entire series of superfamilies (Crowson, 1971) is
represented in the Karatau collection, but it has not been possible to prove
the validity of many identifications.

A few primitive hydrophilids have been found among the Staphylinifor-
mia, including forms close to Hydraenidae. In addition, many staphylinids
(nearly 200 impressions) of the most primitive subfamilies Oxytelinae* and
Tachyporinae have been found; these belong to 16 species and 10 genera
(Tikhomirova, 1968, 1973).

The Eucinetiformia are represented by rather abundant Eucinetidae.
These differ only slightly, at least in external appearance, from the present-
day Eucinetus.

The Scarabaeiformia are rare in Karatau. Beetles externally resembling
Dascilloidea are fairly widespread but their dascilloid nature and the validity
of their inclusion in the genus Mesodascilla (Martynov, 1925) of the Dascil-
lidae, has not been confirmed. Representatives of Scarabaeidae are also less
numerous. Only two rather poorly preserved beetles have been found. These
are close to the primitive Scarabaeidae already abundant in the Early
Cretaceous.

Fossils of the Elateriformia are particularly abundant in Karatau. The
only strictly Mesozoic group of Polyphaga, which could be assigned the
taxonomic status of a superfamily, has been found here. Its members are
rather diverse in morphology and size; several dozen of them have already
been found. These are closest to representatives of the recently established
superfamily Artematopoidea Crowson, 1973, but are distinguished by the
presence of a transverse metasternal suture. This character unites them with
metallic wood borers. The latter are rare in Karatau; the number of specimens

*Given as Oxytelidae in the Russian original—General Editor.

14

13

collected is less than 10. Click beetles are the most abundant group of
Polyphaga, and of beetles in general, in Karatau. Nearly 250 of their
specimens have been collected. They mainly belong to about 20 genera of
the primitive, extinct subfamily Protagrypninae (Dolin, 1975) distributed
almost exclusively in the Mesozoic. Members of the Agrypninae, Diminae,
Negastriinae and Cardiophorinae are rarely encountered. Members of the
Bostrychiformia have not been identified among the Karatau beetles, but are
almost certainly present.

The Cucujiformia are rather well represented in Karatau. The Cleroidea
are few though diverse. The Cucujoidea are represented by primitive
(Parandrexis) as well as advanced (Praemordella and Jurallecula) forms.
The Chrysomeloidea are represented by the rather diverse Protoscelinae,
considered to be phytophages. The total absence of Cerambycidae is
noteworthy. Rhynchophora are particularly abundant in Karatau. They all
belong to the newly established, primitive family Eobelidae which is ex-
clusively Mesozoic. Eobelid fossils are slightly less numerous than click
beetle fossils. They are very diverse; 30 species and 16 genera grouped into
four subfamilies are described in this text.

The Karatau beetles are also ecologically very diverse. The Jurassic
Lake of Karatau apparently had normal* salinity. Absolutely no aquatic
insects lived in it; in any case no fossil larvae have been found. The adult
beetles (schizophoroids, gyrinids, coptoclavids, hydrophilids) fell into the
lake during flight and died. The diversity of Coptoclavidae in Karatau may
be emphasized once again; the number of genera found here is greater than
in all the remaining sites. The Karatau period** apparently coincided with
the maximum flourishing of this strictly Mesozoic, highly specialized family.
The well-known Solenhofen site in the Federal Republic of Germany is
second in the diversity of its coptoclavids; this site is almost coeval with
Karatau. The littoral coastal beetles—to which many, if not all, Geadephaga
and Staphylinidae belong—are fairly abundant in Karatau.

Terrestrial phytophagous beetles from Karatau are mainly associated
with woody plants and are represented by xylophages, more accurately, by
xylomycetophages. Distinctly phyllophagous forms are absent among them.
Even the phytophagous Protoscelinae, judging from the mode of life of their
closest extant relatives, inhabited pachycaulous sago palms (Cycadophyta)
or Bennettitales. A large proportion of the dendrobionts were probably
confined to conifer trunks. In any case, the abundance of cupedids in the

*This is a correct translation of the Russian, but from the context it seems that the word
“abnonnal” may have been intended—Scientific Editor.

**We are not aware of any Karatau period. Since the reference is obviously to the Karatau
site this sentence can be reworded as “The Karatau site apparently exhibits the greatest diversity
of this strictly Mesozoic, highly specialized family”—General Editor.

14

Mesozoic correlates well with the abundance of conifers and with the level
of carbon accumulation in predominantly conifer rich region. Cupedids were
the most widely distributed of the Jurassic dendrobionts. They fed on wood
partly decomposed by fungi. Possibly, only metallic wood borers fed on fresh
wood. The remaining highly abundant Karatau Elateriformia were associated
with wood highly decomposed by fungi. The ecology of the Eobelidae is of
considerable interest. Their larvae probably lived in the ovules and strobilae
of cycadoid gymnosperms, Cycadophyta and Bennettitales. The ecology of
some present-day Oxycorynidae, as well as the correlation between the
abundance of Eobelidae and that of the Cycadophyta, supports this assump-
tion.

Coleopteran fossils from the dawn of the Early Cretaceous (Neocomian)
are distributed rather unevenly. The fossils collected from the Upper Jurassic
Baisa site in the Vitim plateau of Trans-Baikal far outnumber those from
other sites, although over a hundred have already been collected from a single
site in Trans-Baikal and Mongolia. Therefore, an attempt will be made to
characterize the Early Cretaceous Coleoptera from this site, while those from
the remaining sites will be introduced only as supplementary material. The
sites yielding early Cretaceous insects are plentiful in Trans-Baikal, Mon-
golia and China. They are also found in the remaining territory of Siberia in
Kazakhstan, but comparably rich sites have not yet been exposed. Isolated
finds are known from western Europe, North Africa, South America and
Australia.

In general features, the Neocomian beetle fauna is similar to that of the
Late Jurassic. The Archostemata are somewhat less abundant but continue
to play an important role, constituting a few percent of the collected fossils.
Eight species belonging to 6 genera and 2 families of the Archostemata have
been described. In some sites cupedids occur in unusually large numbers or
even predominate. All these sites are poor in fossils, but cupedids turn out to
be over represented. It may simply be that chance has favoured their selection
from a fauna in which they actually constitute only a small percent. Among
the Archostemata, cupedids predominate in all faunas. Of the remaining
families of the Archostemata, the only representatives are the ademosynids.
However, one cannot speak of these beetles with absolute certainty as of the
Archostemata*. Judging from the elytral composition, schizophoroids still
occurred in this period, but not a single well preserved fossil of these forms
has been found. Among cupedids, the Notocupes predominates as usual; the
exclusively Late Mesozoic genera are very rare, and so are the Cenozoic type
of cupedids.

Fossil Adephaga are very abundant in most collections of Early
Cretaceous beetles. This predominance is due to the vast collections of larvae

*Considering what has been said in the first part of this sentence the end part should read
“... as of other members of the Archostemata’”—General Editor.

15

15

of a single species of water beetle, Coptoclava longipoda Ping, which already
exceed a thousand. However, even if we exclude this species, the eee
still numerically exceed the Archostemata. PEATE beY

To date, coptoclavids and whirligigs are the only aquatic Ae gne in
the Lower Cretaceous sites. The examples of Coptoclavidae include: the most
advanced, widespread, monotypic genus Coptoclava, from the Early
Cretaceous of East Asia; and the most primitive Necronectes, represented by
a single fossil from Brezina in North Africa. The whirligigs possess the
typical Mesozoic facies. The Geadephaga are represented by ground beetles
and trachypacheids. Both* genera of trachypacheids already existed in the
Late Jurassic; the carabid genera [from the Lower Cretaceous sites**] were
specific, but for the most part primitive, and not well differentiated from the
Late Jurassic forms. Only one fossil may be related to the present-day
carabids, but it is very incomplete. The discovery of an aquatic larva of
Cretotaenia with the full complement of abdominal segments is noteworthy.

The Polyphaga are diverse and abundant, but possess a primarily
Mesozoic appearance. Among the Staphyliniformia, rather diverse
hydrophilids and staphylinids of the lower} subfamilies are found. Among
the Eucinetiformia, Eucinetidae similar to present-day Eucinetus are en-
countered; they are definitely more abundant than others. The Scarabaeifor-
mia are extensively represented for the first time in the Early Cretaceous. In
the Lower Cretaceous deposits of East Asia, fossil Scarabaeidae are the most
abundant among the terrestrial beetles. Compared to the Late Jurassic, their
numbers rose more than tenfold. Despite their high diversity and excellent
preservation, it is difficult to ascertain the taxonomic position of the Early
Cretaceous Scarabaeidae. Its members are externally similar to the fairly
advanced forms, often resembling May beetles. However the important
structural features of this family, such as the form of the hind wings and
spiracles, when evident, are found to be extremely primitive, and confirm
their inclusion in the lowest Scarabaeidae#, including the Geotrupidae which
are presently considered a separate family.

Elateriformia, abundant in the Late Jurassic, become exceedingly rare
in the Early Cretaceous. The metallic wood borers increase slightly in
number, whereas the click beetles become distinctly fewer. The Protagryp-
ninae lose their dominance among the click beetles, and the Oestodinae make
their appearance (V.G. Dolin, personal communication). Also noteworthy is
the first appearance of the Throscidae and Cerophytidae.

*Since names of these genera have not been given above it is not certain that the author
implied two instead of ‘both’—General Editor.

**_Scientific Editor.

+More primitive—Scientific Editor.

+So given in the Russian original. Should read Scarabaeidae (sensu lato)—General Editor.

16

The Cucujiformia were previously not as important as now. The
Cucujoidea are represented by the Nitidulidae and Scraptiidae, essentially
indistinguishable from present-day forms. The Chrysomelidae are and ap-
parently: still represented only by Protoscelinae. In the Baisa site, beetles
externally similar to the primitive Cerambycidae have been found, but their
incomplete preservation precludes their exact placement.

The Early Cretaceous Rhynchophora are diverse; even primitive
belidoids are present which, according to L.V. Amol’di may be directly
related to Belidae and Oxycorynidae, but V.V. Zherikhin considers that they
belong with Eobelidae. Also, representatives of the Attelabidae and Cur-
culionidae are found for the first time. Ecologically, the Early Cretaceous
beetles preserve the Mesozoic character. The aquatic beetles are represented
by pleustonic predators (adult Coptoclavidae and Gyrinidae); nektonic
predators (larvae of Coptoclava); and hydrophilids, which are decidedly
phytophagous in the adult stage. The latter, despite their general resemblance
to Spercheus, were not adapted to the pleustonic mode of existence, and
probably fed mainly on charophytic algae while moving among submerged
vegetation.

Terrestrial predators are represented by staphylinids, carabids and
trachypacheids of Mesozoic appearance, which were evidently dwelling near
water. It is even possible that the aquatic caraboid larvae of Cretotaenia
belonged to the trachypacheids.

Among the abundant Scarabaeidae, probably there were no forms
similar in mode of life to the modern, cosmopolitan dung beetles and May
beetles; instead, they were probably mycetophages and carrion-feeders. A
large proportion of the beetles in the Early Cretaceous as well as the Late
Jurassic were xylobiontic phytophages associated with wood more or less
decomposed by fungi. As usual, phyllophages were few or absent among the
beetles. Among weevils there are forms obviously associated with
angiosperms; they are still very rare, but their remains have been found in
these deposits (Bakhrameev, 1973).

The insect composition of the Early Cretaceous is quite unique. An
extensive proliferation of angiosperms occurred during this period and,
apparently linked with this, a change in the composition of the entomofauna.
The majority of the Late Mesozoic forms characteristic of the end of the
Jurassic and beginning of the Cretaceous disappear; they were quite unex-
pectedly replaced, not by Cenozoic forms, but by the Late Mesozoic forms
not found earlier in taphocenoses and considered totally extinct. Unfortunate-
ly, we do not have rich, well-preserved fossils of large beetles from the Aptian
and Albian stages. They are either small or fragmentary and cannot be
accurately identified. In the Trans-Baikal sites, whose age was determined
as the end of the Early Cretaceous, fossils of Liadytidae have been found
which were previously encountered only in the first half of the Jurassic.

17

Trachypacheidae and Hydrophilidae very near the Early Jurassic forms were
also found at these sites. Finally, from the Middle Cretaceous deposits of the
Labrador Peninsula, a beetle has been found which was described as a
member of the monotypic family Labradorocoleidae. This family may be
related exclusively to beetles of the family Tshekardocoleidae which are
characteristic of the Early Permian and, even in the Late Permian, so far only
one specimen has been found (Ponomarenko, 1969a). Based on elytral
composition, this site resembles the Early Jurassic; schizophoroid type elytra,
not observed in younger deposits, predominate. Apparently, relict forms
occurred in the new biocenoses; the forms typical of relatively stable
Mesozoic and Cenozoic biocenoses began to disappear while new relatively
stable cenoses with typical Cenozoic groups had not yet fully formed.

The insects of the Late Cretaceous can be studied far better than those
of any other period. We have not only sites with the imprints of the Late
Cretaceous insects but also numerous sites with insect remains embedded in
fossil resins, providing a good sequence of forms through the entire Late
Cretaceous. The study of these collections has just begun and it is possible
to present only the first tentative results.

Cenozoic forms predominate among the Late Cretaceous insects, just as
angiosperms do among the Late Cretaceous plants. The Mesophyte-
Cenophyte boundary also delineates one of the greatest changes in the
evolution of beetles. The composition of the Late Cretaceous beetle fauna is
essentially Cenozoic. It is true that the abundance of Archostemata, par-
ticularly cupedids, does not decrease; in Siberia their numbers even increase.
This may have been linked to the extinction of chekans, while conifers
retained their importance. In the late Cretaceous, cupedids were the only
Archostemata, represented by the very widespread Mesozoic genus
Notocupes and the rarely encountered examples of present-day Omma and
Tetraphalerus.

Strictly Mesozoic forms are totally absent from the Late Cretaceous
Adephaga. Dytiscidae, Gyrinidae and Carabidae of the present-day type are
found here, while advanced forms such as Harpalinae* are already present
in the first half of the Late Cretaceous. Most of the subfamilies and tribes of
extant Adephaga had apparently already evolved in the Late Cretaceous.

A similar picture is seen for the Late Cretaceous Polyphaga which have
been almost totally excluded from the investigation. All these are closely
related to the Cenozoic forms; strictly Mesozoic groups are either completely
absent (Eobelidae and Protoscelinae) or are very rare (Protagrypninae).
Unfortunately, it is not yet known whether phyllophages were present among
the Late Cretaceous beetles, but Cenozoic longhorned beetles and metallic
wood borers were undoubtedly more common than in the Early Cretaceous.

*Family Carabidae—Scientific Editor.

18

In conclusion, it may be said that two major faunal changes occurred in
the evolution of Coleoptera during the Mesozoic: the first between the Early
and Late Jurassic when the Archostemata lost their dominance, and the
second at the end of the Early Cretaceous when the Mesozoic fauna was
replaced by the Cenozoic.

17

Descriptions of New Taxa

[In order to present measurements in a format which will be more familiar
to most of our readers, Russian expressions such as “‘A twice shorter than B”
have been translated as either “A half as long as B” or “B twice as long as A”.
Decimals have been transformed into fractions when convenient—Scientific
Editor. ]

SUBORDER ADEPHAGA
Family TRIAPLIDAE Ponomarenko, Fam. Nov.

Diagnosis. Fairly large, elongate, cylindrical beetles with head bent ventral-
ly. Head small, ortho- or opisthognathous. Antennae weakly moniliform.
Prothorax with distinct sternopleural and notopleural sutures. Forecoxae
protuberant, not separated by intercoxal process. Mesosternum short,
mesopleuron small. Suture dividing mesothoracic postepisternum distinct.
Middle coxae converging. Metasternum roundly tapering anteriorly; mete-
pisternum extending to middle coxal cavities; transverse metasternal suture
distinct. Hind coxae fully separating metasternum and abdomen, long, with
large femoral plates covering basal abdominal sternites. Abdomen with six
visible sternites, the first small, and last noticeably longer than the remaining.
Elytra without punctate striae, with a projection situated laterally on inner
side*.

Composition. Single genus in the Triassic of Soviet Central Asia.

Comparison. Differs from other families, except Haliplidae, in the
structure of hind coxae and abdominal base; differs from Haliplidae in
ventrally bent head, contiguous forecoxae without intercoxal process,
metepisterna extending to middle coxal cavities, and shorter hind coxae with
smaller femoral plates.

*The relevant figures suggest the following interpretation of the preceding phrase: “with a
projection [= ridge or shelf] situated /aterally [= toward the costal margin] on inner side [= the
inflexed portion of elytron, the elytral epipleuron].”—Scientific Editor.

18

20
Genus Triaplus * Ponomarenko, gen. nov.

Species name coined from Triassic, and genus Haliplus.

Type species. T. macroplatus sp. nov.; Triassic of Soviet Central Asia.

Description. Body elongate, teardrop-shaped, posteriorly tapering.
Head short, strongly transverse, retracted under pronotum, often entirely
concealed. Antennae fairly long, slender: basal segments cylindrical, distal
markedly moniliform. Pronotum transverse, forming semi-circular hood
above the head. Prosternum rectangular, shorter than forecoxae. Propleura
not at all narrowing anteriorly. Mesosternum much shorter than middle
coxae. Middle coxae rounded. Metasternum transverse, strongly and roundly
tapering anteriorly, its hind margin and transverse suture nearly straight.
Hind coxae slightly shorter than broad, laterally deeply incised with the
portion extending backward along the midline almost at right angles to the
coxal base. Femoral plates approximately as long as wide. Abdomen marked-
ly longer than the meso- and metasternum together, last sternite much longer
than the penultimate. Elytra long, extended, pointed apically with distinct
inner projection** approximately level with hind coxae. Body with large and
dense punctures.

Species composition. Two species in the Triassic of Soviet Central
Asia.

Triaplus macroplatus Ponomarenko, sp. nov.
(Plate I, Photos 1, 2; Figure 1)

Species name coined from ‘macros’ (Greek)—large, and ‘plata’ (Greek) —
plate.

Holotype. No. 2971/104, PIN, impression of beetle without antennae,
legs and abdomen, Soviet Central Asia, Southern Fergana, Kirgiz SSR, Osh
Oblast, Batken region, Madygen area (Madygen site, south-western part).
Triassic, Madygen series.

Material. Besides holotype, three impressions of beetles, Nos. 2083/
163, 2905/24 and 2971/111, from the same site.

Description. Prosternum slightly shorter than forecoxae; propleuron
narrow, elongate. Forecoxae equal in length and width, mesally protuberant.
Mesosternum one-third length of middle coxae. Middle coxae somewhat
transverse. Metepisternum strongly broadened anteriorly. Femoral plates
together forming an almost perfect semicircle. True abdominal sternites
II-VI equal in iength, half the length of the last. Legs short, femora
uniformly thickened, tibiae widening from base to apex. Inter-puncture

* Given as Triaplidae in the Russian original—General Editor.
** See note on page 19—Scientific Editor.

21

distance on dorsal surface of body nearly equal to (diameter of) punctures
themselves, punctures much smaller on ventral surface. Punctures on all
ventral sclerites identical.

Dimensions. Body length about 7-8 mm, width 2.9-3.3 mm, elytral
length 6.5—7.0 mm.

Comparison. Distinguished by longer forecoxae and shape of femoral
plates together forming an almost perfect semicircle.

Triaplus laticoxa Ponomarenko, sp. nov.
(Plate I, Photo 3; Figure 2)

Species name coined from ‘latus’ (Latin}—wide, and ‘coxa’ (Latin)—pelvis.

Holotype. No. 2240/179, PIN, impression of almost entire beetle, shape

of the impression slightly obliterated by postfossilization expansion of the

19 rock. Soviet Central Asia, Southern Fergana, Kirgiz SSR, Osh Oblast, Batken

region, Madygen area (Dzhailyaucho site, northern part). Triassic, Madygen
series.

18 Fig. 1. Triaplus macroplatus sp. nov. a, b—holotype PIN No. 2971/104: a—dor-
sal view; b—ventral view; c—paratype PIN No. 2905/24. Madygen, Triassic. Here
and in the following figures the linear scale is approximately 1 mm.

De

Material. Besides holotype, seven almost entire impressions of beetles,
Nos. 2069/1228, 2069/1294, 2070/1887, 2240/216, 2240/218, 2240/243,
2555/1720, from the same site. Fossils of beetles rather highly variable in
size and possibly belong to more than one species. However, structurally
very similar, but a comparison of the proportions of individual sclerites is
very difficult due to obliteration associated with expansion of the embedding
rock.

Description. Length of head half its width. Prosternum slightly shorter
than forecoxae, propleuron of subequal length and width. Forecoxae some-
what transverse. Mesosternum half the length of middle coxae, middle coxae
equal in length and width. Metepisternum strongly broadening anteriorly,
narrowing at posterior margin. Femoral plates somewhat broadening from
base posteriorly, and rounded only just before apex. Abdominal sternites
IJI-VI equal in length, half the length of the last, the seventh. Legs short,
hind femora slightly clavate, broadest in basal third. Body dorsally with
tubercles and uniform punctures, ventrally with large dense punctation,
punctures on metasternum and femoral plates particularly large.

Fig.2. Triaplus laticoxa sp. nov. a, b—holotype PIN No. 2240/179: a—dorsal view,
b—ventral view of head and thorax; c—paratype PIN No. 2555/1720. Dzhailyaucho,
Triassic.

20

2)

Dimensions. Body length 11-14 mm, width 3.54.0 mm, elytral length
10.0-11.5 mm.

Comparison. Distinguished by longer prosternum, and femoral plates
broadening posteriorly from base, extending along the middle coxae*.

INFRAORDER HYDRADEPHAGA

SUPERFAMILY DYTISCOIDEA LEACH, 1815
Family PARAHYGROBIIDAE Ponomarenko, Fam. Nov.

Diagnosis. Water beetles. Larvae nektonic, with natatorial legs and urogom-
phi. Head sessile, rectangular, with roundish projecting nasale. Antennae
four segmented, nearly as long as head. Mandibles short, strongly curved,
with pointed elongate apices and wide base, mesal surface with small
denticles (retinaculae). Prothorax somewhat longer than other body segments
which are each of nearly equal length. Abdomen eight segmented; last tergite
somewhat prolonged posteriorly at middle, overlapping the only functional
pair of spiracles. All 3 pairs of legs similar, bearing sparse stuff bristles.
Urogomphi long, distinctly segmented, with apical bristles.

Composition. Single genus from the Jurassic of Trans-Baikal.

Comparison. Most closely resembles the Hygrobiidae in the structure
of head and legs, but differs from it in the structure of mandibles and retention
of the eighth abdominal segment; it differs from most dytiscoids by the
presence of retinaculum, and from the forms with retinaculum by the shape
of head and natatorial legs, long urogomphi, and short eighth abdominal
segment; from the Coptoclavidae it differs in the shape of mandibles and in
possessing stiffer and sparser bristles on uniformly wide leg segments.

Remarks. Description of the new family is based almost exclusively on
plesiomorphic characters of water beetles. Thus affinity of Parahygrobia
with any family of water beetles is based on symplesiomorphy. Parahygrobia
does not have features which allow it to be considered an ancestor of water
beetles; it is highly unlikely that such benthic phytophages as the Noteridae
originated from them.

Genus Parahygrobia Ponomarenko, gen. nov.

Species name coined from the genus Hygrobia.
Type species. P. natans sp. nov.; Jurassic of Trans-Baikal.
Description. Small larva, robust with rather long legs and urogomphi.
Head almost square, slightly broadening anteriorly. Nasale small, length

* As given in the Russian original, but presumably the hind coxae are the structures
indicated here—Scientific Editor.

21

24

much less than width, its demarcation distinct basally at middle. “Epicranial
suture” Y -shaped, its stem longer than fork. Length of antennal segments less
than double the width. Pronotum narrowed anteriorly. Eighth abdominal
segment slightly shorter than the preceding; tip of seventh tergite angularly
projecting posteriorly. Urogomphi slightly shorter than the abdomen,
noticeably narrowing apically, urogomphal hairs longer than the segments.
Legs long; femora and tibiae slightly thickened, tarsi virgate; femora, tibiae
and tarsi with long natatory bristles, their length slightly less than that of the
distal leg segments. Claws large, their bases broad, not narrower than the tip
of tibia, only half the length of tarsus.
Species composition. Monotypic genus.

/

Fig.3. Parahygrobia natans sp. nov. Holotype PIN No. 3053/423: a—ventral
view, b—dorsal view of head and prothorax, c—tip of abdomen. Uda, Jurassic.

a

21

25

Parahygrobia natans Ponomarenko, sp. nov.
(Plate I, Photo 4; Figure 3)

Species name coined from ‘natans’ (Latin)—natatory.

Holotype. No. 3053/423, PIN, impression of entire larva. Trans-Baikal,
Buryat ASSR, Eravninsk region, River Uda close to Ulan-Mailo settlement
(Uda site): Jurassic, Uda series.

Description. Central part of nasale occupying nearly one-third its width,
apically rounded (Fig. 3b). Epicranial suture bifurcated midlength of the
head, its branches initially diverge, then run parallel almost to the anterior
margin of the head, near which they again sharply diverge laterally. Lateral
ocelli on a single sclerite*, separated by more than their diameter. Antennae
slightly shorter than the head, first segment shorter, fourth longer than the
subequal second and third segments. Mandibles slightly longer than wide,
retinaculum blunt and small in the apical third. Three proximal segments of
maxillary palps nearly equal, the terminal segment noticeably thinner than
the proximal ones.

Pronotum almost equal to the head in length and to its width anteriorly,
anterior pronotal margin nearly half the width of posterior margin. Meso-
and metathorax each two-thirds the length of prothorax. All abdominal
segments of nearly equal length, eighth slightly shorter than the remaining
segments. Width of the widest (fourth abdominal) segment twice the width
of eighth at base. Urogomphi three-fourths the length of abdomen, distinctly
10-segmented. Legs only slightly shorter than body. Coxae large, oval, with
distinct longitudinal ridge, slightly shorter than femur. Trochanters single,
one-fourth the length of femur. Forefemora uniformly convex at middle,
one-and-a-half times longer than tibiae and tarsi, each bearing three bristles
on the posterior margin. Foretibiae with a pre-apical notch, claws of foretarsi
Straight, slightly shorter than tarsi. Bristles on foretibiae and tarsi denser on
the posterior margin than on the anterior margin. Middle tibiae and femora
equal, slightly shorter than tarsi; claws half as long as tarsi. Femora with a
few bristles borne preapically on the posterior margin. Tibiae and tarsi with
5-7 bristles on the anterior and posterior margins. Hind femora and tibiae
longer than middle ones, equal to each other, distinctly shorter than tarsi.
[Tarsal] claws unequal: larger claw half the tarsal length, straight; shorter
claw thicker and curved.

Dimensions. Length of larva 5.8-6.5 mm, length of head 0.8-1.0 mm,
length of foretarsus 0.4-0.5 mm, midtarsus 0.5—0.6 mm, length of urogomphi
1.4-1.5 mm.

Remarks. Sediments with Parahygrobia larvae are considered Upper
Jurassic (Skoblo, 1968), but some larvae of aquatic insects collected here

*The Russian original uses the term ‘apodeme’ which has been changed to
sclerite—Technical Editor.

22

26

cannot be distinguished from those of the Middle Jurassic on the basis of the
described characters. Hence the problem of their precise age remains un-
resolved (for greater details see Rasnitsyn, 1975).

Family COPTOCLAVIDAE Ponomarenko, 1961

Coptoclavidae Ponomarenko, 1961, p. 68; Rohdendorf and Ponomarenko,
1962, p. 225; Ponomarenko, 1975, p. 123.

Diagnosis. Large, rarely medium-sized, aquatic beetles. Head with two
pairs of eyes, dorsal and ventral. Metepisternum not reaching middle coxae,
hind coxae not broadening anteriorly, transverse metasternal suture absent.
Middle and hind legs natatory, tibiae broadened, together with tarsi forming
an oar-like lobe.

Larvae nektonic, actively natatory, with cylindrical body not tapering
posteriorly, and with long urogomphi. Mandibles with denticles on mesal
surface. Forelegs raptorial, with spines; middle and hind legs natatory, with
thickened tibiae and tarsi, bearing slender and dense natatory hairs. Abdomen
8-segmented, spiracles metapneustic, urogomphi with fused segments.

Composition. Three subfamilies: Necronectinae in the Jurassic of
Europe and Asia, and in the Early Cretaceous of Africa; Charonoscaphinae
in the Jurassic of Asia, and Coptoclavinae in the Early Cretaceous of East
Asia.

Comparison. Distinguished from all natatory beetles as a four-eyed
form with natatory middle and hind legs, and hind coxae not anteriorly
broadening; from whirligigs distinguished by long legs and dominance of
metathorax over mesothorax.

Subfamily Necronectinae Ponomarenko, Subfam. Nov.

Diagnosis. Large beetles. Hind coxae with small femoral plates. Hind tibiae
and tarsi long and slender without any broadening, and with long, slender,
natatory hairs.

Composition. Genus Necronectes found in the Middle Jurassic of Trans-
Baikal, Late Jurassic of South Kazakhstan and Early Cretaceous of Algeria;
genus Exedia in the Late Jurassic of South Kazakhstan; genus Pseudo-
hydrophilus in the Late Jurassic of Western Europe. Aquatic larva of
Stygeonectes from the Middle and possibly Late Jurassic Baikal is tentatively
placed here.

Comparison. Distinguished from the remaining subfamilies by slender
hind tibiae and tarsi in adults and by femoral plates on hind coxae. The
Stygeonectes larvae are distinguished from the Coptoclava larvae by the
single denticle on the mesal surface of the mandible and by the proportions
of leg segments.

23

2
Genus Necronectes Ponomarenko, gen. nov.

Species name coined from ‘necros’ (Greek)—dead, and ‘nectes’ (Greek)—
oarsman.

Type species. N. aquaticus sp. nov. Middle Jurassic of Baikal.

Description. Large beetles with rather elongate body expanded at
shoulders. Head and pronotum relatively small. Head transverse, strongly
retracted into the anterior emargination of the prothorax. Pronotum expanded
in front of posterior angles. Mesothorax only slightly smaller than
metathorax, metapleura larger than mesopleura. Metepisterna broadening
slightly anteriorly their inner margins almost without a curve; anteriorly not
reaching middle coxal cavities. Transverse metasternal suture faint or not
noticeable. Femoral plates scarcely longer than and about three times as wide
as hind coxae, strongly tapered laterally. Elytra smooth. All legs with
natatory hairs. Fore- and middle tarsi broadening, their proximal segment
long, broadening apically, second and third transverse, notched apically, last
segment virgate.

Species composition. N. aquaticus from the Middle Jurassic of Baikal
(Ust’-Balei), and N. giganteus from the Upper Jurassic of south Kazakhstan
(Karatau). The incomplete fossils from the Early Cretaceous of Algeria and
the Jurassic of Soviet Central Asia are included in this genus.

Comparison. Distinguished from other genera by basally expanded
pronotum.

Necronectes aquaticus Ponomarenko, sp. nov.
(Plate I, Photo 5; Figure 4)

Species name coined from ‘aquaticus’ (Latin)—aquatic.

Holotype. No. 722/23, PIN, almost entire impression beetle. Baikal,
Irkutsk, Oblast, Irkutsk region, right bank of river Angara, below Ust’-Balei
village (Ust’-Balei site). Lower strata of the Middle Jurassic, Cheremkhovsk
series.

Material. Besides the holotype, one poorly preserved impression of a
male and part of an isolated elytron, specimen Nos. 2363/15 and 2363/17
from Khudoga site (Buryat ASSR, Dzhidin region, Dzhida river basin,
waterdivide of Khudoga and Simkhak rivers, Ichetui series). The impression
of the beetle differs somewhat from the holotype in the shape of body and of
fore femora, but resembles the holotype in most other characters.

Description. Body convex, oval, flattened dorsally, with medium keel
ventrally. Head and pronotum together small, approximately one-sixth the
body length. Length of head almost half the occipital width, narrowing
anteriorly. Dorsal eyes small, almost one-fourth the length of head, separated
by more than their diameter. Ventral eyes on each side of head, situated

28

anterior to dorsal ones, elongate. Gular plate short and wide, almost square,
mental sections almost equal. Antennae slender, filiform, longer than the
head and prothorax together, segments cylindrical, only scape noticeably
broadening apically, two apical segments shorter than the rest.

Length of prothorax one-third its width, widest in the anterior third, its
lateral margin with a depression in front of posterior angles. Pronotal angles
rounded, anterior margin not emarginate, concealing the base of the head
dorsally. Pleura* wide, narrowing anteriorly. Prostemum projecting roof-
like**, shorter than forecoxae and prosternal process. Mesosternum small,
bordering middle anteriorly, laterally strongly tapered, with shallow notch
in middle. Mesopleuron almost square, twice as long as mesosternum, suture
between mesepisternum and mesepimeron dividing the pleuron into equal
triangular parts. Mesepimeron extending to middle coxal cavities only as
narrow tongue. Metathorax transverse; width of metasternum more than
twice the distance between middle and hind coxae, its lateral margins almost
straight, hind margin angularly projecting backward. Transverse metasternal
suture bent backward, weak. Longitudinal metasternal lines, corresponding
to internal ridges, run from middle coxae backward and from hind coxae
forward. Metepisternum triangular, its anterior margin half the length; [much
longer than+] distance between inner corners of metepisternum and middle
coxae much less than the length of anterior margin of metepisternum. Length
of hind coxae with femoral plates half the width. Portion of hind coxae
projecting over the abdomen almost one-third length of entire coxa. Femoral
plates transverse, sharply tapered in lateral third.

Abdominal sternites subequal in length; only the last (seventh) slightly
longer and in males with a shallow, wide, apical notch [exposing an addi-
tional sternite]. Eightht sternite [of male] paired, irregular oval, with a
flattened medial side and notched antero-lateral margin. Spiracles of seventh
and eighth segments each one-third the length of seventh sternite, almost
twice as large-as the preceding spiracles. Parameres}} almost symmetrical,
triangular, only slightly longer than wide.

Elytra smooth, epipleuron narrow, a long ridge present near lateral
[= costal?] margin at middle third of elytron on lower surface. Forecoxae
slightly projecting, almost round, length of trochanter nearly one-third length

*The nature of description suggests the reference to the propleuron. If accepted, ‘pleura’

should be changed to ‘propleura’—General Editor.

**Presumably the transverse anterior portion for the prostemmum, minus its intercoxal process
is referred to here—Scientific Editor.

+As in the Russian original but with brackets added. Deleting the bracketed phrase will
restore meaning—Scientific Editor.

+As in the Russsian original but contradicts the earlier statement—General Editor. This
contradiction is eliminated with the addition of the bracketed interpolative remarks—Scientific
Editor.

++Given as parameters in the Russian original—General Editor.

29

: an

24 Fig. 4. Necronectes aquaticus sp. nov. Holotype PIN No. 722/23: a—ventral view,
b—dorsal view, c—ventral view of head and thorax, d—antenna, e, f—forelegs, g,
h—midlegs, i—hind legs.*

* Only one hind leg shown—General Editor.

24

25

30

of femur. Tibia slightly shorter than femur, flattened and curved, slightly
broadening apically. Tarsi subequal in length, length of the first segment
almost equal to the next three; second and third broadened, with apical notch,
third segment articulating with the notch of second; fourth segment small,
almost square, fitting into notch of third; fifth twice as long as the preceding.
Spur of foretibia shorter than first tarsal segment.

Middle femora one-third longer than forefemora. Tibiae two-thirds
length of femora, flattened, apically broadening, their lateral margin serrate.
Middle tarsus three-fourths the length of tibia; its first segment only slightly
shorter than remaining segment together, twice as long as its apical width;
second and third transverse, third with deep apical notch; fourth and fifth
narrow, fourth a little longer, fifth twice as long as wide. Claws two-thirds
the length of apical tarsal segment. Tibia and tarsus with median groove
bearing long and dense natatory hairs.

Hind trochanters one-third the length of femora. Femora uniformly
slightly thickened, hind femora as long as middle femora. Tibiae markedly
longer than femora, broadening apically and somewhat curving externally.
Tarsi noticeably shorter than tibiae, their segments elongate, slightly
broadening apically; first tarsal segment equal in length to three apical
segments together, three times longer than second; second and third segments
equal; fourth two-thirds of third; fifth almost equal to two preceding ones
together.

Body with dense shallow punctures dorsally, almost forming transverse
striae on elytra; smooth ventrally except for a few rather small punctures on
the sternum. First and second visible abdominal sternites with large circular
punctures laterally.

Dimensions. Body length 34-35 mm, width 15 mm, elytral length
25 mm.

Comparison. Distinguished by the very small head and pronotum, long,
drawn-out body, and fine punctures on elytra.

Necronectes gigas Ponomarenko, sp. nov.
(Plate I, Photo 6, Figure 5)

Species name coined from ‘gigas’ (Latin)—gigantic.

Holotype. No. 2554/477, PIN, beetle with incompletely preserved legs.
South Kazakhstan, Chimkent Oblast, Alagabass region, south-western flank
of Kashkarata river valley, Aulie area in Mikhailovka village (Karatau-Mik-
hailovka site); Upper Jurassic, Karabastau series.

Description. Large beetle with oval, flattened body. Head and pronotum
together about one-fourth of body length. Length of head two-thirds its width.
Dorsal eyes rather large, round, distance between ocular sclerites* less than

*See our note on page 25—General Editor.

26

31

their diameter. Ventral eyes round, placed laterally on head in front of dorsal
ones. Antenna short, filiform, not reaching the base of pronotum, its segments
noticeably broadening apically; scape cylindrical, short, third segmenta little
longer than others.

Length of prothorax half its maximum width just before middle.
Pronotum narrowed anteriorly, roundly emarginate at anterior margin, ex-
panded in front of posterior angles. Mesosternum small, with distinct depres-
sion for prosternal process. Mesopleura almost square, pleural suture
dividing them into equal triangular parts. Metasternum four times wider than
distance between middle and hind coxae. Metepisternum rather wide
posteriorly, anteriorly wider than at posterior margin, almost double; middle
coxae separated from inner corners of metepisternum by half the length of
latter at anterior margin. Length of hind coxae two-fifths the width; portion
of coxa projecting over abdomen longer than wide, its length half the width
of entire coxa. Femoral plates almost square, two-fifths the coxal width,

=

Fig.5. Necronectes gigas sp. nov. Holotype PIN No. 2554/477: a—ventral view,
b—dorsal view. Karatau, Upper Jurassic.

32

sharply tapering laterally, their notches almost rectangular. Abdomen
posteriorly strongly narrowing, almost pointed at apex, last sternite tn-
angular, first visible and third to fifth sternites equal in length, last longer but
second longest.

Elytra smooth, epipleuron narrow, a long ndge present near lateral
[= costal?] margin at middle third of elytron. Forefemora thickened; tibiae
much shorter than femora and uniformly broadening to apex. Tarsi equal in
length to tibiae. First tarsal segment triangular, longer than the three follow-
ing together; second to fourth transverse, slightly broadened apically, weakly
notched at apex. Last segment slightly narrower than preceding segments.
Middle tibiae longer than foretibiae, two-thirds the length of middle femora,
tarsus slightly shorter than tibia, first segment slightly broadening from base
to apex, scarcely longer than the three following segments together; second
to fourth segments transverse, weakly notched apically. Last segment slightly
longer than preceding segment. Spur of middle tibia not extending to apex
of first tarsal segment. Hind femora not longer than middle femora, equal to
tibiae. Hind tibiae slightly broadening apically, flattened, with a groove
bearing natatory hairs. Tarsus noticeably shorter than tibia, its first segment
one-and-a-half times longer than the subequal second and third segments.
Tarsal segments slender, cylindrical.

Body dorsally with dense punctures, finer on head and pronotum,
coarser on elytra. Large punctures of elytra fused into intermingling
transverse striae. Punctures small and sparse ventrally. First abdominal
sternite with large circular punctures on each side of femoral plates.

Dimensions. Body length 48 mm, width 23 mm; elytral length 33 mm.

Comparison. Distinguished from N. aguaticus sp. nov. by large head
and pronotum and posteriorly strongly tapering abdomen, from N. cyrenaicus
sp. nov., by narrow, laterally highly tapering femoral plates and from N.
planus sp. nov. by transverse hind coxae.

Material. Holotype.

Necronectes cyrenaicus Ponomarenko, sp. nov.
(Plate I, Photo 7; Figure 6)

Species name coined from ‘Cyrenaica’ (Latin)—Roman province in North
Africa.

Holotype. No. 3394/24, PIN, metathorax, abdomen and elytra of beetle.
North Africa, Algeria, Saida department, 20 km from Brezina settlement.
Lower Cretaceous, Goteriv*? intermediate sandstone stratum.

*Only here and on page 82 Goteriv is spelt with double ‘t’ elsewhere it is with one ‘t’,
hence the preference—General Editor.

i es i il

27

27

313)

Fig. 6. Necronectes cyrenaicus, sp.nov. Holotype PIN No. 3394/24: a—ventral view
of abdomen and tibia*, b—apices of elytra and folded wings. Algeria, Brezina, Lower
Cretaceous.

Description. Large, oval, flattened beetle. Inner face of elytra in anterior
third, with along projection next to lateral margin. Epipleuron narrow. [Hind]
wing with “oblongum” cell narrow, its medial width two-fifths its length.
Length of hind coxae half their width; portion of coxa projecting over
abdomen two-fifths width of entire coxa. Femoral plate transverse, tapering
gradually in lateral third, lateral notch rounded, shallow. Second abdominal
sternite one-and-a-half times longer than third. Abdomen strongly narrowing
posteriorly to pointed apex. Elytra with coarse, large, sparse punctures, fusing
into transverse striae and forming longitudinal rows along tracheae. Ab-
dominal sternites with fine, frequently contiguous punctures; punctures
noticeably larger laterally on first visible and second abdominal sternites.

Dimensions. Body length about 45 mm, width 24 mm, elytral length
about 35 mm.

Material. Holotype.

Comparison. Distinguished from N. aquaticus sp. nov. by posteriorly
strongly tapering body, and from N. gigas sp. nov. by broad femoral plates.

Necronectes latus Ponomarenko, sp. nov.
(Plate I, Photo 8; Figure 7)

Species name coined from ‘latus’ (Latin)—wide or broad.

*Tibia is seen to occupy the position of femur—General Editor.

28

34

Holotype. No. 3073/151, PIN impression of the thorax, abdomen and
elytra of beetle. Soviet Central Asia, Kirghiz SSR, Batken region, Sagul
Ravine (Shurab-III site). Jurassic.

Material. Holotype.

Description. Mesosternum very short, mesopleuron almost square,
mesepisternum and mesepimeron rectangular, equal in length. Mesepimera
wide, forming a part of the wall of middle coxal cavities. Metasternum almost
three times wider than distance between rounded middle and hind coxae,
strongly narrowing anteriorly, pesterior margin angularly projecting back-
ward. Transverse metasternal suture not visible. Metepisternum anteriorly
broadening; width of anterior margin almost one-third the length. Length of
hind coxae with femoral plates noticeably less than width; femoral plates
strongly tapering laterally, transformed beyond middle of coxae into a
narrow tongue running along the anterior coxal margin but not quite reaching
its lateral corners. Last abdominal sternite twice as long as the remaining
equal sternites, its width two-thirds the width of abdominal base, apex blunt,
rounded. All spiracles almost uniform in size. Elytra broad, flattened, sharply
narrowing pre-apically. Epipleuron narrow. Integument of body very thin,
wings and abdominal spiracles visible through it. Only coxal apices and
margins of the last three abdominal sternites are sclerotized and pigmented.
Elytra almost smooth, body with fine and sparse punctation ventrally, dis-
tance between punctures larger than the punctures themselves.

Dimensions. Body length about 15 mm, width 6.4 mm; elytral length
11.5 mm.

Comparison. Differs sharply from other species of the genus by broad
abdominal apex, highly oblique hind coxae and laterally sharply tapering
femoral plates. Probably merits separation into an independent genus, but its
imperfect preservation prevents its full comparison.

Genus Exedia Ponomarenko, gen. nov.

Genus name coined from ‘exedia’ (Greek)—float.

Type species. E. plana. Upper Jurassic of South Kazakhstan.

Description. Large beetles with flat, oval body. Head large, transverse,
dorsal eyes convergent. Pronotum rather strongly, roundly tapering anterior-
ly, emarginate in front. Metepisterna almost reaching middle coxal cavities.
Posterior abdominal sternites shorter than preceding [anterior] ones.
Forecoxae short, tibiae shorter than femora. Hind tibiae flattened, narrow,
slightly curved. Elytra with raised ridges.

Special composition. Monotypic genus.

Comparison. Distinguished from other gnera by ridged elytra; in addi-
tion, differs from Necronectes in having pronotum not expanded at base and
from Pseudohydrophilus in the shape of foretarsi.

35

Fig. 7. ? Necronectes latus, sp. nov. Holotype PIN No. 3073/151. Shurab-Ill.
Jurassic.

Exedia plana Ponomarenko, sp. nov.
(Plate II, Photo 1; Figure 8)

Species name coined from ‘plana’ (Greek)—flat.

Holotype. No. 25554/478*, PIN, almost entire impression of beetle,
without antennae, fore- and middle tarsi. South Kazakhstan, Chimkent
oblast, Algabass region, south-western flank of Kashkarata river valley,
Aulie area in Mikhailovka village (Karatau-Mikhailovka site). Upper Juras-
sic, Karabastau series.

Material. Holotype. In addition, paired elytra of a larger beetle found at
the same site, specimen PIN, No. 2239/1322 (elytral length 50 mm, width 11
mm), very similar in shape. Due to large difference in dimensions, its positive
inclusion in this species is problematic.

Description. Length of head nearly half the width, deeply retracted into
emargination of anterior pronotal margin. Pronotal length half its width at
base, its anterior margin half the width of posterior. Prosternum visible in
front of forecoxae, prosternal process projecting anteriorly as a flat keel.
Mesosternum shorter than large middle coxae, mesepimera strongly
broadening towards shoulders. Metasternum roundly tapering anteriorly,
length nearly one-third the width; its anterior margin half the width of
posterior. Distance between middle and hind coxae greater than length of

* Given as No. 2554/778 both in Fig. 8 and Plate II, Photo 1—General Editor.

29

29

36

hind coxae. Length of metepisterna much greater than its width at anterior
margin. Width of hind coxae 1.7 times the length, central raised part twice
as long as wide. Abdomen slightly broadening to base of second visible
sternite and tapering thereafter. First sternite slightly shorter than subequal
second and third; fourth sternite two-thirds the length of third and equal to
sixth; fifth sternite shorter than them. Elytra with alternate higher and lower
ridges.

Legs rather long. Foretibiae noticeably broadening apically. Middle
femora thickening medially, nind femora weakly clavate, broadening in
apical third. Hind tibiae equal to femora in length, but much narrower. Hind
tarsi much shorter than tibiae. Integument highly sclerotized, with fine
punctures.

Dimensions. Body length 32 mm, width 18 mm; elytral length 23 mm.

Genus Pseudohydrophilus Deichmiiller, 1886

Pseudohydrophilus: Deichmiiller, 1886, p. 67; Handlirsch 1906-1908, p.
544; Ponomarenko, 1971, p. 75. Prodytiscus: Oppenheim., 1888, p. 237.

ly U \
(Sep
AQ a
od

xN \ / 4

1 ee
ES
SH

b

Fig. 8. Exedia plana, sp. nov. Holotype PIN No. 2554/778; a—dorsal view,
b—ventral view. Karatau, Upper Jurassic.

30

37

Poor preservation of earlier material of this genus precludes exact
determination of its taxonomic position. However, definitive chracters do not
contradict its inclusion in the Coptoclavidae whereas, all chracters con-
sidered together, it cannot be related to any other known taxon. In one
specimen, femoral plates chracteristic of Necronectidae may be observed
(Ponomarenko, 1971a, Plate VIII, Photo 2). This genus differs from the
representatives of the subfamily [Necronectinae] in the structure of the
foretarsi, the middle segments of which are not notched apically.

Genus Stygeonectes Ponomarenko, gen. nov.

Genus name coined from the mythical River Styx (Greek), and ‘nectes’
(Greek)—oarsman.

Type species. S. jurassicus sp. nov. Jurassic of Baikal.

Description. Rather large, cylindrical, natatory larvae with long, oar-
shaped middle and hind legs. Head sessile, weakly transverse, widest in front
of posterior angles, anterior margin nearly linear, nasale absent. Lateral ocelli
six, rather large, on a single sclerite* near anterior angles of head capsule.
Antennae almost equal to head in length, first and fourth antennal segments
much shorter than second and third. Mandibles shorter than their apical
width; apical denticles pointed, directed medially. Retinaculum pointed,
situated approximately at middle of mandible. Proximal to retinaculum,
mandible much wider than distal to it.

Prothoracic tergite longer than meso- and metatergites; first seven
abdominal tergites shorter than thoracic and the last abdominal tergite.
Femora, tibiae and tarsi of forelegs subequal in length, with short and fine
bristles. Middle and hind femora noticeably shorter than tibiae and tarsi.
Tibiae and tarsi with dense, long and slender natatory hairs on both sides.
Upper margin of tarsi convex. Eighth abdominal tergite transverse; its
posterior margin scarcely overhanging the only open spiracles, located below
the base of the urogomphi at middle of hind margin. Well-developed tracheae
proceed anteriorly from the spiracles. Urogomphi long, rather thick, straight
with indistinct segmentation, lacking setae.

Species composition. Monotypic genus.

Remarks. Small, early-stage larvae of Stygeonectes closely resemble
Parahygrobia larvae. They differ in the shape of the mandibles, absence of
urogomphal pads and in the fine and dense natatory hairs. However, all these
structural features can be noticed in only a very few of the fossil remains.
Therefore, the best distinguishing feature is the shape of the middle and hind
tarsi. In Parahygrobia they are virgate, their upper margin (usually the
anterior margin in the impressions) is straight. In Stygeonectes the anterior

*See our note on page 25—General Editor.

38

margin of the tarsi is curved, roundly protuberant. Stygeonectes displays this
unique character while swimming*, Coptoclava lavae similarly use the
middle and hind pairs of legs for swimming. This similarity forces us to
consider Stygeonectes and Coptoclava as members of one group, whereas
Parahygrobia remains among the commen natatory forms.

Stygeonectes jurassicus Ponomarenko, sp. nov.
(Plate II, Photos 2, 3; Figure 9)

Species name coined from ‘jurassicus’ (Latin)—Jurassic.

Holotype. No. 3000/935, PIN, almost entire middle-instar larva. Trans-
Baikal, Buryat ASSR, Mukhorshibir region, Novospasskoe settlement
(Novospasskoe site). Middle Jurassic, Ichetui series.

Material. Seventy larvae, of which SO in collection No. 3000 from
Novospasskoe site; 4 in collection No. 3053 from Uda site, Buryat ASSR,
Eravnin region, Uda river close to Ulan-Mailo (tribal) settlement [village],
Udin series; 2 in coliection No. 3436 from Borzhe site, Buryat ASSR,
Eravnin region, Borzhin depression, well No. 35, depth 154-156 m, Udin
series; 4 in collection No. 2363 from Khudoga site, Buryat ASSR, Dzhida
region, Dzhida river basin, watershed of Khudoga and Simkhak rivers,
Ichetui series; 6 in collection Nos. 1980, 1981, 1982 from Ichetui site, Buryat
ASSR, Dzhida region, upper reaches of Ichetui river at Petropavlovka
settlement, Ichetui series; 2 in collection No. 2089 from Bukukun site, Chita
oblast, Kyrin region, left bank of Bukukun river, Bukukun series; 1 in
collection No. 1873 from Ust’-Balei site, Irkutsk oblast, Irkutsk region,
Angara river near Ust-Balei settlement, Cheremkhov series; 1 in collection
No. 1669 from lya site, Irkutsk oblast, Tulun region, left bank of river Iya
near Vladimirovka village, Cheremkhov series.

Description. Head (Fig. 9, b, d) of late-instar larva almost rectangular,
of early-instar noticeably narrowing anteriorly. Head capsule with Y-shaped
ecdysial line on vertex (“epicranial suture’’), its common stem almost equal
in length to the fork. Lateral ocelli ovoid, rather large, distance between them
smaller than their diameter. First antennal segment equal in thickness to
second segment, but one-fourth its length, third thicker and slightly shorter
than second, fourth segment two-thirds as thick as third, scarcely longer than
first.

Body of larva hardly narrowing posteriorly, flattened dorsoventrally. Its
lower part not sclerotized, well-defined sclerites present only at the base
of legs. Prothoracic tergite of late-instar larvae approximately one-and-a-half

* Probably a misprint in the original. Meaning can be restored by replacing “displays” with
“uses” or by substituting “for” in place of “while’—Scientific Editor.

By)

Fig. 9. Stygeonectes jurassicus, sp. nov. a, b—holotype PIN No, 3000/935:

a—dorsal view, b—head, c—paratype PIN No. 3000/975, head, d —paratype No.

3000/939, e—paratype PIN No. 3000/986; all from Novospasskoe, Jurassic,

f—paratype PIN No. 3053/426; Uda, Jurassic; g—paratype PIN No. 1873/11;
Ust’-Balei, Jurassic.

40

times longer than the subequal meso-and metathoracic tergites and almost
equal to the eighth abdominal tergite. In early-instar larvae all the thoracic
tergites and the last abdominal tergite are subequal, longer than the remaining
abdominal tergites.

Forecoxae nearly half the length of forefemora, tibiae and tarsi. Middle
and hind coxae longer than forecoxae, slightly shorter than the equal distal
segments of legs. Middle tarsi longer than foretarsi, claws of foretarsi much
longer than claws of middle and hind tarsi. Posterior margin of middle and
hind femora and both margins of tibiae and tarsi with fine natatory hairs, their
length exceeding width of the segment. On tarsi, natatory hairs more than
ten. Urogomphi more than three times as long as last abdominal segment.

32 Measurements Larval instars
I II Ill

Body without urogomphi 4.8 7-15 21-24
Prothoracic tergite 0.4-0.7 1.0-1.1 1.5-1.8
Mesothoracic tergite 0.3-0.5 0.8-1.0 1.2-1.4
Eighth abdominal tergite 0.4-0.5 1.0-1.2 —
Foretarsi 0.5—0.7 0.8-1.5 Po)
Middle and hind tarsi 0.7-0.8 1.0-1.7 2.3-2.5
Urogomphi 1.5-2.2 2.7-3.2 —

Measurements show considerable variations in the size of S. jurassicus
larvae, much more than in Coptoclava longipoda larvae. Besides the total
length varying mainly with the degree of gaseous inflation of the larvae
during postmortem putrefaction, the size of the sclerites also varies. How-
ever, no regular trends in this variability could be detected. Dimensions vary
widely not only in larvae from different sites but also in those from the same
Novospasskoe site, where the majority of them were collected. There is wide
variation in the size of sclerites of middle-instar larvae. An impression is
created that these sclerites are of larvae of two different instars. However,
statistical reliability of these differences could not be established on the
available material; for now it would be expedient to say that Stygeonectes
had three larval instars, as is common among beetles.

Ecology. Predatory, actively swimming Stygeonectes larvae inhabited
small, sparsely vegetated and generally flowing lakes of Western Trans-
Baikal and Baikal. The raptorial forelegs and cutting mandibles, very similar
to the mandibles of ground beetles of the genus Carabus, show that these
larvae, like those of Coptoclava longipoda, caught their prey with forelegs
and then tore it apart. Late-instar larvae are very rarely encountered.

41

Remarks. Fossil remains of Stygeonectes larvae are found in deposits
assigned various ages. The Cheremkhov series generally relate to lower
Dogger, Ichetui to its upper beds, and the Udin series to lower Malm (Skoblo,
1968). As mentioned earlier, large variability and insufficient material do not
permit detection of either reliable differences among larvae collected from
various sites or their exact identity.

Subfamily Charonoscaphinae Ponomarenko, Subfam. Nov.

Diagnosis. Large and medium-sized beetles. Hind coxae without femoral
plates. Hind tibiae broad and flat; tarsi flat but not broad, much narrower than
tibiae. Hind tibiae and tarsi with long natatory hairs.
Composition. Two genera in the Late Jurassic of South Kazakhstan.
Comparison. Differs from the Necronectinae in the absence of femoral
plates and expanded hind tibiae, and from the Coptoclavinae in the narrow
hind tarsi and natatory hairs.

Genus Charonoscapha Ponomarenko, gen. nov.

Genus name coined from Charon of Greek mythology and ‘scapha’
(Greek)—boat.

Type species. C. grossa, sp. nov. Upper Jurassic of South Kazakhstan.

Description. Large and medium-sized beetles with flattened body. Head
small, but transverse. Dorsal eyes well spaced, ventral eyes situated in front
of them on both sides of head. Pronotum strongly narrowing anteriorly, in
front of deep emargination. Metepisterna not reaching middle coxal cavities.
Terminal abdominal sternites shorter than preceding ones. Forecoxae rather
long, tibiae slightly longer than femora, first and last segments of foretarsi
elongate, but each not longer than three middle segments together. Middle
and hind tibiae flat and broad, with a fringe of long and slender natatory hairs.
Middle tarsal segments flat and broad in males, but flat and linear in females.
Hind tarsi with long, flat and linear segments and natatory hairs. Elytra with
protuberant ridges in males, almost smooth in females, with interweaving
rows of punctures.

Species composition. Two species in the Late Jurassic of South
Kazakhstan.

Comparison. Distinguished by anteriorly deeply emarginate pronotum.

Charonoscapha grossa Ponomarenko, sp. nov.
(Plate II, Photos 4, 5; Figure 10)

Species name coined from ‘grossum’ (Latin)—large.

42

Holotype. No. 2554/446, PIN, almost entire impression of male. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Mikhailovka site).
Upper Jurassic, Karabastau series.

Material. Besides the holotype, one female No. 2997/1847 from the
same site.

Description. A large beetle with broad parallel-sided body. Length of
head two-thirds its width, deeply retracted into the emarginate anterior

d

Fig. 10. Charonoscapha grossa, sp. nov. a, b—holotype PIN No. 2554/446:
a—ventral view; b—dorsal view of posterior end of body; c, d—paratype PIN No.
2997/1847 : c—ventral view; d —head and pronotum. Karatau, Upper Jurassic.

43

34 margin of pronotum. Antennae slender, filiform, scarcely extending beyond
the pronotal base, first segment noticeably larger than the remaining, second
short, equal in length and width, remaining segments elongate, last short.
Pronotum strongly roundly tapering anteriorly, its anterior margin half the
width of posterior, maximum length two-fifths the width. Prosternum
anterior to forecoxae shorter than coxae. Propleura wider than long. Proster-
nal process broadening toward apex. Mesosternum small, shorter than middle
coxae, anteriorly with longitudinal groove for the prosternal process.
Mesepisterna subequal in length and width. Mesepimera narrow and long,
noticeably broadening toward shoulders. Middle small, weakly protuberant,
posteriorly covered by metasternal process. Metasternum much shorter than
wide, strongly narrowing anteriorly, distance between hind coxae shorter
than between middle coxae. Metepisterna short, strongly broadening
anteriorly. Hind coxae triangular, short; length about two-thirds the width.
Medial raised part of coxa almost three times longer than its width. Abdomen
narrowing posteriorly from second visible sternite; last sternite small, short,
its length one-third the width. Elytra flattened, epipleura strongly broadening
anteriorly. Ridges on elytra rather weak in male, homonomous.

Legs short, foretibiae almost not flattened toward apex, weakly curved;
first tarsal segment in male scarcely wider than tibia and one-third its length;
second to fourth segments noticeably broadening apically, transverse, very
slightly wider than first segment, fifth equal to the first in length, narrow,
tapering apically. Tibial spur long, reaching the apex of fourth tarsal segment.
Claws short. Middle femora broad, longer than forefemora. Middle tibiae
shorter than femur, curved and broadening toward the apex. Middle tarsus a
little longer than tibia, its first segment two-fifths the length of tibia, in males
its length is twice the width, in females four times the width. Spur of middle
tibia not reaching the apex of first tarsal segment. Hind femora approximately
equal in size to middle femora, tibiae shorter than them, broadening and
flattening toward apex. Tarsi flat, with linear segments. Valvulae sclerotized.
Integument with fine, dense punctation.

Dimensions. Body length 24-26 mm, width 12-13 mm; elytral length
18-19 mm.

Comparison. Distinguished by parallel-sided body, strongly anteriorly
narrowing pronotum and short last abdominal sternite.

Charonoscapha ovata Ponomarenko, gen. nov.
(Plate II, Photos 6, 7; Figure 11)

Species name coined from ‘ovaius’ (Latin)—oval.

Holotype. No. 2997/1845, PIN, impression of female without head,
abdomen, fore- and middle legs. South Kazakhstan, Chimkent oblast,
Algabass region, south-western flank of Kashkarata river valley, Aulie area.

35

44

Mikhailovka village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau
series.

Material. Besides holotype, three incomplete remains, one male and
two females, specimen Nos. 2066/2393, 2784/1925, 2997/404, from the
same site, and one female specimen No. 2452/439 from the Karatau-Galkino
site.

Description. Medium-sized beetles with flat, ovoid, posteriorly strong-
ly tapering body. Head together with prothorax forming an almost perfect
semicircle. Length of head somewhat less than width, labrum rounded.
Antennal segments rather short and thick. Anterior margin of pronotum half
the width of posterior, pronotal width 2.3 times the maximum length; anterior
margin of pronotum with a deep emargination. Prosternum slightly shorter
than forecoxae; prosternal process not broadening posteriorly, propleura
slightly longer than wide. Mesosternum short, almost taenioid [ribbon-
shaped]. Metepisterna noticeably wider than long; mesepimera strongly
broadening toward shoulders. Middle coxae small, slightly projecting;
metasternal process between them narrow. Metasternum short, but distance
between the middle and hind coxae greater than length of middle coxae.
Metepisterna broad, strongly broadening anteriorly. Hind coxae triangular,
short, their length nearly half the width; median raised part of coxa twice as
long as wide, posteriorly broadening, its outer angle drawn out. Abdomen
roundly tapering; beginning with second visible sternite, all sternites of equal
length; length of the last sternite two-fifths its width. Elytra of male with
distinct, well defined ridges, the four main ridges twice as high as inter-
mediate ones. Elytra of female with numerous dense rows of punctures. Body
with fine punctation. Forefemora somewhat dilated, tibiae apically broaden-
ing, tarsal segments in female slender, linear. Middle femora longer than
forefemora; [the corresponding] tibiae shorter than them, broadening apical-
ly, shorter than tarsi. Tarsi linear, with flattened segments. Hind femora
shorter than middle femora, slightly broadening in the proximal half. Tibiae
slightly curved and flatter in one-third*, noticeably broader than femora and
flat; segments of tarsi about half as wide as tibiae.

Dimensions. Body length 15.6-17.9 mm, width 7.5-8.4 mm, elytral
length 11.5-12.8 mm.

Comparison. Distinguished by oval, posteriorly narrowing body, and
longer last abdominal sternite.

Genus Charonoscaphidia Ponomarenko, gen. nov. Genus name coined
from genus Charonoscapha.

Type species. C. elongata, Upper Jurassic of South Kazakhstan.

Description. Rather large, elongate-oval beetles. Head triangular, al-
most as long as wide. Dorsal eyes contiguous. Pronotum trapezoidal, nar-

*Jt is not clear from either the description or the relevant figure which third of the tibiae are
so modified—Scienufic Editor.

36

BD

45

rowing anteriorly, not emarginate anteriorly. Metepisterna not extending to
middle coxal cavities. Forecoxae rather long, projecting. Hind tibiae broad
and flat, with natatory hairs. Elytra smooth.

Species composition. Monotypic genus.

Comparison. Distinguished from genus Charonoscapha gen. nov. by a
longer head, trapezoidal pronotum without an emargination on the anterior
margin.

Charonoscaphidia elongata Ponomarenko, sp. nov.
(Plate II, Photo 8; Figure 12)

Species name coined from ‘elongata’ (Latin)—elongated.

Holotype. No. 2384/717, PIN, impression of beetle (female ?) without
fore- and middle legs. South Kazakhstan, Chimkent oblast, Algabass region,
south-western slope of Kashkarata river valley, Aulie area near Mikhailovka
village (Mikhailovka site). Upper Jurassic, Karabastau series.

Fig. 11. Charonoscapha ovata, sp. nov. a —holotype PIN No. 2992/1845, ventral
view; b—paratype PIN No. 2066/2393, dorsal view. Karatau, Upper Jurassic.

46

Material. Besides holotype, two incomplete impressions of beetles,
specimen Nos. 2384/707, 2384/718, and paired incomplete elytra, specimen
No. 2066/2591 from the same site. Almost entire impression of beetle,
specimen No. 2465/904 from Karatau-Galkino site.

Description. Head triangular, slightly longer than wide, almost not
retracted into prothorax. Pronotum not longer than head, anterior margin
two-thirds the width of posterior; maximum length half the width of posterior
margin. Prosternum shorter than forecoxae; propleura longer than wide.
Forecoxae rather long, projecting. Hind coxae triangular, only slightly
shorter than wide, their raised middle part three times longer than wide.
Abdomen narrowing posteriorly from base, length of the last sternite about
half its width. Hind tibiae same length as femora but a little wider than them.
Elytra with distinct dense rows of punctures. Body with fine punctation.

Dimensions. Body length 23 mm, width 10 mm; elytral length 16—
17 mm.

Fig. 12. Charonoscaphidia elongata sp. nov., holotype PIN No. 2384/717:
a—dorsal view; b—ventral view. Karatau, Upper Jurassic.

en

37

Fig. 13. Coptoclava longipoda Ping, reconstructed. a —dorsal view, b—ventral
view; c—larva. East Asia, Lower Cretaceous.

Bil,

38

48
Subfamily Coptoclavinae Ponomarenko, 1961

Diagnosis. Large beetles. Hind coxae without femoral plates. Middle and
hind tarsi flat and broad, not narrower than tibiae, without natatory hairs.
Larvae with mandibles having two denticles on mesal surface, foretarsi much
longer than tibiae, middle and hind tarsi broad; urogomphi highly sclerotized,
slightly curved.

Composition. Single genus Coptoclava Ping (Fig. 13) in Early (Neoco-
mian) Cretaceous of Trans-Baikal, Mongolia, North and East China.

Comparison. Adult beetles distinguished from members of the remain-
ing subfamilies by homonomous structure of middle and hind legs with
distinctly broad tarsi lacking natatory hairs and by mandibles having
numerous denticles; larvae are distinguished by the proportions of legs.

Family LIADYTIDAE Ponomarenko, Fam. Nov.

Diagnosis. Rather small aquatic beetles with oval, biconvex body. Head
noticeably retracted into the prothorax, scutellum external. Metepisterna*
reaching middle coxal cavities. Metasternum with median longitudinal
protuberance, sharply delimited laterally. Hind coxae transverse, not
broadening anteriorly, their anterior margin almost straight with small central
emargination, bounded in front by an extension of the transverse metasternal
suture which forms a single straight line with the anterior margin of the coxae.
Legs slender and long. Hind tibiae not shorter than femora and tarsi, very
slender, linear. Tarsal segments not broad or oar-shaped. Tibiae and tarsi with
natatory hairs.

Composition. One genus in the Mesozoic of Asia.

Comparison. Distinguished from all natatory forms by the very long
and slender hind legs, particularly tibiae and tarsi. Differs from Dytiscidae
in the presence of a metasternal suture and by the not-anteriorly broadening
hind coxae. Differs from Noteridae and Coptoclavidae in the presence of a
metasternal suture and metepisterna extending to middle coxal cavities.
Differs from Hygrobiidae by the metepisterna extending to middle coxal
cavities and by the flat, protuberant area on the metasternum, and from
Amphizoidae by the natatory hind legs and flat protuberant area on the
metasternum.

Remarks. Liadytidae are plesiomorphic in almost all characters used for
distinguishing the families of natatory forms. Only the flat protuberant area
on metasternum, uniting Liadytidae and Noteridae, may be considered
synapomorphic; but based on the structure of the hind legs, these families
occupy extreme positions among the natatory forms—in Liadytidae the legs

*Given as ‘Mesepistema’ in the Russian original, but subsequently given as “metepisterna’
in the comparisons with other families—Scientific Editor.

39

49

are longest, and in Noteridae shortest. In the geochronological distribution
of Liadytidae, two natatory larvae of highly generalized structure—
Parahygrobia and Angaragabus—were found. The latter even occurred in
the same site as one of the species of Liadytes. Either of them could be the
larva of Liadytes, but the difference between them far exceeds the usual
interspecific difference between larvae of the same natatory genus [therefore
itis highly unlikely that both Parahygrobia and Angaragabus are synonyms
of Liadytes—Scientific Editor].

Genus Liadytes Ponomarenko, 1963
Liadytes: Ponomarenko, 1963, p. 129.

Liadytes longus Ponomarenko, sp. nov.
(Plate III, Photo 1, Figure 14)

Species name coined from ‘longus’ (Latin)—long
Holotype. No. 2046/1, PIN, reverse impression of beetle without mid-
dle and hind legs. Trans-Baikal, Chitinskaya Oblast, Shelopuginsk region,

Sve il

oP

Fig. 14. Liadytes longus, sp. nov.; holotype PIN No. 2046/1: a—dorsal view;
b—ventral view. Daya, Lower Cretaceous.

39

50

left bank of Daya river, 4.5 km north of Daya settlement (Daya site; Lower
Cretaceous, Aptian-Albian, Balei series).

Material. Holotype.

Description. Highly elongate, oval, medium-sized beetles. Length of
body exceeds maximum width at posterior third by two-and-a-half times.
Head small, two-thirds the width of occiput, tapering straight from base.
Genae and temples very short, less than half the diameter of eyes. Antennae
weakly moniliform, extending to pronotal base; first segment equal to the
following two together; second slightly shorter than third; remaining anten-
nal segments subequal, slightly longer than wide. Gular plate wide, anteriorly
broadening, occupying about one-third width of head on posterior margin.
Lateral lobes of submentum much larger than median lobe.

Prothorax slightly shorter than head, anteriorly strongly narrowing,
anterior margin two-thirds width of posterior. Length of prosternum two-
fifths the width at posterior margin. Mesosternum long, not shorter than
prosternum. Metasternum transverse, very strongly narrowing anteriorly;
anterior margin one-third the width of posterior. Width at posterior margin
2.7 times the length. Posterior extension of metasternum separated by
transverse suture, rounded. Metepisterna very strongly broadening anterior-
ly, their length only slightly more than anterior width. Hind coxae transverse,
their lateral length almost half that along midline of body; width 2.7 times
the length. Anterior margin of coxa slightly projecting forward from the side.
Elevated middle part of coxae posteriorly broadening, rounded at posterior
margin; its length two-and-a-half times the width. Length of abdomen nearly
half the body length, narrowing posteriorly only from the base of third
sternite. First abdominal sternite two-thirds the length of second which is
equal to the third and fourth together; fifth very slightly shorter than the
fourth; sixth equal to the third, longer than the fourth. Length of sixth sternite
two-fifths the width. Middle and hind femora uniformly thickened, three
times broader than tibiae. Hind femora, tibiae and tarsi equal in length.
Segments of hind tarsi two-thirds the width of tibiae, almost of equal length.
Natatory hairs slender, apical spines not visible on segments of hind tarsi.
Integument of body with fine, dense punctation.

Dimensions. Body length 9.2 mm, width 3.7 mm; elytrallength 7.0mm.

Comparison. Distinguished by elongate body, siightly antero-laterally
broadening hind coxae, and rounded posteromedial extension of meta-
sternum.

Liadytes crassus Ponomarenko, sp. nov.
(Plate III, Photo 2, Figure 15)

Species name coined from “crassus” (Latin)—broad.

40

51

Holotype. No. 3015/369, PIN, impression of metathorax, base of ab-
domen and hind legs. Trans-Baikal, Chitinskaya Oblast, Balei region, right
bank of Unda river, 2 km above Zhidka settlement (Unda site). Lower
Cretaceous, Aptian-Albian? Balei series.

Material. Holotype.

Description. Oval, medium sized beetle. Length of metasternum one-
fourth the width at posterior margin, anterior margin half the width of
posterior; posterior margin angularly extending backward in middle third.
Longitudinal and transverse metasternal sutures distinct. Hind coxae
transverse, only slightly tapering laterally. Their lateral length two-thirds the
length along midline of body, maximum length half the width. Anterior coxal
margin projecting anteriorly. Elevated middle part of coxa slight tapered
posteriorly; posteriorly with lateral notch, apically truncate; length of
elevated part more than half its width. Abdomen narrowing posteriorly from
third sternite. First abdominal sternite two-fifths the length of second, third
half that of second. Hind femur, tibia and tarsus approximately equal in
length. Hind femur markedly broadening along almost its entire length, only
three times as long as wide. Hind tibia one-fourth the width of hind femur.
Three basal segments of hind tarsus equal in length. Spurs of hind tibiae and
spines on apices of tarsal segments stiff. Integument of body with rather large
distinct punctation.

Dimensions. Body length about 6 mm, width 3.0 mm, elytral length
4.3 mm.

Fig. 15. Liadytes crassus, sp. nov.; holotype PIN No. 3015/369. Unda, Lower
Cretaceous.

4

—y

52

Comparison. Resembles L. longus, in the anteriorly extending hind
coxae but differs in the posterior triangular extension of metasternum, thicker
hind femora, and wider body.

Family DYTISCIDAE Leach, 1815
Genus Cretodytes Ponomarenko, gen. nov.

Genus name coined from “creta” (Latin)—chalk, and genus Dytiscus.

Type species. C. latipes; Upper Cretaceous, Turon of South Kaza-
khstan.

Diagnosis. Lateral extensions of metepisterna [metapleurosternites]
narrow, triangular; apical recess on anterior projection of metasternum
triangular, deep and rather wide. Elevated middle plates of hind coxae
slightly diverging posteriorly, not broadening toward apex, apically rounded.
Hind legs broad; tibia two-thirds the length of femur, approximately two-
and-a-half times as long as wide. Preapical groove and associated tuft of hairs
lacking on hind femur. Width of femur at its articulation with the trochanter
not narrower than the trochanter.

Species composition. Monotypic genus.

Taxonomic position. A single fossil of thorax and hind legs has been
found, a formal comparison of the genus is therefore not possible. Cretodytes
must belong either to the subfamily Colymbetinae or to the Dytiscinae based
on the following characters: metapleurosternites rather wide; middle plate of
coxa distinctly notched before its anterior margin, lateral notch absent and
apex [= hind margin] rounded. Within the first [= Colymbetinae], based on
the broad metapleurosternites and absence of the characteristic hair tuft of
the femoral apices, Cretodytes would have to be placed in the tribe Colym-
betini; the structure of the anterior metasternal extension is also similar.
However, such broad and short tibiae are not found in the Colymbetini,
moreover the middle plates of the hind coxae are usually broadening towards
the apex and strongly divergent in this tribe. Within the Dytiscinae,
Cretodytes resembles Dytiscini in possessing the fairly broad metapleuro-
sternites and in the position of the middle plates of the hind coxae, but in the
Dytiscini, these plates are almost never rounded apically, and such small
beetles are unknown. Thus, by indirect characters [evidence], Cretodytes
cannot be definitely included in any one present-day natatory tribe since it
shares features of the most primitive tribes of both Colymbetinae and
Dytiscinae. It is hardly advisable, however, to describe a suprageneric taxon
from such an incomplete fossil.

3)

Cretodytes latipes Ponomarenko, sp. nov.
(Plate III, Photo 3, Figure 16)

Holotype. No. 2383/256, PIN, impression of metasternum and incom-
plete hind legs. South Kazakhstan Kyzyl-Ordin Oblast, Chiilii region, north-
western spurs of Karatau range (Kyzyl-Dzhar site). Upper Cretaceous,
Turon, Beleutin series.

Material. Holotype.

Description. Rather small beetles. Metasternum strongly transverse,
depression on its anterior extension triangular; length of metasternum one-
and-a-half times its width, middle part raised and flattened, with narrow
median longitudinal furrow. Metapleurosternites triangular, their length be-
tween points of greatest proximity with middle and hind coxae is half the
distance from the lateral corners of middle coxae to the lateral corners of
metapleurosternites. Medial length of metasternum three times the minimum
intercoxal distance. Hind coxae longest somewhat lateral to their middle, and
laterally tapered beyond. Raised middle part of coxa negligibly narrowing at
about midlength; its width one-fourth the length; apical thirds of coxal plates
diverge, forming an acute-angled triangle. Hind femora approximately four
times longer than wide, three times longer than trochanters. [Hind] tibia not
narrower than femur, with longitudinal rows of marginal spines, two spines
closer to anterior margin.

Dimensions. Length of fossil remains 4.5 mm, width 6.0 mm; length of
beetle thus slightly more than 10 mm and width about 7 mm.

Remarks. Alongside the fossil lay a leg with a very long femur and tibia
(Fig. 16b). By its size it could be the middle leg of the same beetle, but
dytiscids with such long middle tibiae are not known.

Fig. 16. Cretodytes latipes, sp. nov.; holotype PIN No. 2383/256: a—metasternum;
b—midleg of probably the same specimen. Kyzyl-Dzhar, Upper Cretaceous.

42

54

Family GYRINIDAE Latreille, 1840
Genus Avitortor Ponomarenko, gen. nov.

Genus name coined from ‘avus’ (Latin)—ancestor, and ‘tortere’ (Latin)—to
whirl.

Type species. A. primitivus,sp.nov. Lower Cretaceous of Trans-Baikal.

Description. Small, flat whirligigs. Head large, highly transverse, ap-
proximately equal to prothorax in length. Eyes small, round, ventral eyes
antero-lateral to dorsal eyes. Pronotum transverse, anterior margin strongly
emarginate. Anterior margin of prosternum straight, prosternal intercoxal
process very small. Scutellum exposed, triangular, rather large. Middle coxae
oval, almost contiguous, oblique, their length two-thirds the width. Metaster-
num not shorter than hind coxae. Hind coxae transverse, triangular, their
length less than width. Sutures between basal abdominal sternites straight,
distinct. Elytra with fine grooves.

Species composition. Monotypic genus.

Comparison. Distinguished from all whirligigs by the long metaster-
num and narrow middle coxae, and from the present-day forms by the short,
transverse hind coxae.

Fig. 17. Avitortor primitivus, sp. nov.; holotype PIN No. 3064/856: a—dorsal view;
b—ventral view. Baisa, Lower Cretaceous.

4

uw

55

Avitortor primitivus Ponomarenko, sp. nov.
(Plate III, Photo 4, Figure 17)

Species name coined from ‘primitivus’ (Latin)—primitive.

Holotype. No. 3064/856, PIN, impression of beetle without legs. Trans-
Baikal, Buryat ASSR, Eravninsk region, left bank of Vitim river below the
mouth of Baisa river (Baisa site). Lower Cretaceous, Valanzhin-Goteriv,
Zazin series.

Material. Holotype.

Description. Body broadly oval, parallel-sided. Width of head 1.7 times
the length, broadest at eyes, slightly narrowing posteriorly from eyes. Dorsal
eyes slightly more proximate than vertical*, separated by three times their
diameter. Length of pronotum approximately one-third the width, noticeably
narrowing anteriorly; anterior emargination of prosternum rectangular, its
depth scarcely less than the length of prosternum in front of forecoxae.
Middle coxae oblique, irregularly oval, slightly more strongly tapering
externally. Length of metasternum two-fifths the width. Distance between
middle and hind coxae greater than the length of the latter. Length of hind
coxae nearly half their width, lateral margin half the length along median
plane. Basal abdominal sternites short, sutures between them distinct, ster-
nites apparently articulate. Elytra with very fine grooves, without traces of
punctures.

Dimensions. Body length 10.2 mm, width 5.0 mm, elytral length 6.1
mm.

Genus Mesodineutes Ponomarenko, gen. nov.

Genus name coined from Mesozoic and genus Dineutes.

Type species. M. amurensis, sp. nov. Upper Cretaceous, Tsagayan of
Amur oblast, Arkhara.

Diagnosis. Medium-sized whirligigs, with broad flattened, posteriorly
tapering body. Mesosternum transverse, its length half the width, only
slightly longer than middle coxae. Middle coxae set apart. Scutellum visible,
triangular. Metapleurosternites laterally elongate, their length less than their
width or medial length of metasternum. Transverse metasternal suture dis-
tinct, almost straight. Hind coxae transverse, their length about half the width.
Lateral length of coxa only slightly exceeds its length along longitudinal
body axis. Posterior margin of coxa with a deep depression. Abdomen very
slightly longer than meso- and metathorax together, uniformly tapering
posteriorly from base of third sternite; base of seventh sternite narrower than
apex of sixth. First sternite short, suture separating it from second indistinct,

*So given in the Russian original; obviously a misprint should read “ventral”—General
Editor.

44

43

56

second and third longer than fourth to sixth and shorter than seventh. Apex
of seventh rounded. Elytra smooth, margined along suture, laterally with
wide, anteriorly broadened flange; apex of elytra rounded.

Species composition. Monotypic genus.

Comparison. Distinguished from present-day whirligigs possessing
smooth elytra by the transverse short hind coxae, from Angarogyrus by
transverse mesosternum and from Miodineutes by short and broad abdomen.

Mesodineutes amurensis Ponomarenko, sp. nov.
(Plate III, Photos 5, 6; Figure 18)

Holotype. No. 2055/74, PIN, beetle without head, prothorax and legs. Amur
oblast, quarry at Arkhara station (Arkhara site). Upper Cretaceous, Tsagayan.

Material. Holotype and isolated elytra, specimens No. 2055/79,
2055/100 from the same site.

Description. Length of beetle exceeds its width by not more than one-
and-a-half times, body roundly tapering posteriorly from shoulders. Length
of mesosternum two-fifths the width; raised medial part of mesosternum
almost triangular; median longitudinal groove proceeding along it posterior-
ly. Middle coxae slightly tapering laterally, their length about half the width.
Intercoxal distance two-fifths the length of middle coxae; distance between

Fig. 18. Mesodineutes amurensis, sp. nov.; a—holotype PIN No. 2055/74,
b—paratype PIN No. 2055/79. Arkhara, Upper Cretaceous.

Sf

lateral corners of coxae two-thirds the distance between apex of middle coxa
and lateral corner of hind coxa. Hind coxae laterally only very slightly shorter
than along median line of body. Width of elevated part of coxa a little less
than its length, coxa strongly tapering to lateral third and broadening toward
lateral margin. Second abdominal sternite equal in length to third, fourth to
sixth equal in length, seventh longer than fifth and sixth together. Apex of
seventh sternite with fringe of short stiff hairs. Margins of elytral disk with
scarcely noticeable rows of punctures. Elytral surface with very small dense
punctation, almost smooth; body ventrally with sparse rather large punctures.

Dimensions. Body length about 11 mm, width 8.5 mm; elytral length
8.0-8.2 mm.

Genus Angarogyrus Ponomarenko, gen. nov.

Genus name coined from Angara*, and ‘gyros’ (Greek)—spinning.

Type species. A. minimus, sp. nov. Middle Jurassic of Cis-Baikal.

Description. Small whirligigs with elongate-oval body. Head large,
strongly transverse, longer than pronotum. Eyes round, dorsal eyes behind
ventral ones and somewhat more closely grouped. Mesosternum long, almost
square, much longer than metastemum. Metasternum short, metapleuroster-
nites forming laterally elongate triangles. Distance between middle and hind
coxae half the length of metasternum. Hind coxae markedly oblique, their
length half the width, their length at lateral margin half that at the median
line of body. Abdomen rather long, uniformly roundly tapering directly from
base. Elytra smooth, apically rounded.

Species composition. Monotypic genus.

Comparison. Distinguished from present-day whirligigs possessing
smooth elytra by short hind coxae, from the genus Mesodineutes by the long —
mesosternum and less proximate middle and hind coxae, from the genus
Miodineutes by the broader abdominal apex, and by apically non-truncated
elytra.

Angarogyrus minimus Ponomarenko, sp. nov.
(Plate III, Photo 7, Figure 19)

Species name coined from ‘minimus’ (Latin)—small.

Holotype. No. 1874/30, PIN, impression of beetle without legs. Cis-
Baikal, Irkutsk oblast, Tulun region, left bank of Iyariver near Vladimirovka
(Iya site). Middle Jurassic, Cheremkhov series.

Material. Holotype.

*Given as Angarida in the Russian original, possibly referring to the Angara River Basin—
General Editor.

58

45 Fig. 19. Angarogyrus minimus, sp. nov.; holotype PIN No. 1874/30. lya, Jurassic.

Description. Body widest in front of middle, tapering posteriorly some-
what more strongly than anteriorly. Dorsal and ventral eyes approximately
the same size, considerably overlapping*. Distance between dorsal eyes
three times their diameter; distance between ventral eyes and posterior
margin of head, half their diameter. Pronotum strongly roundly tapering
anteriorly, its length less than one-third the width at posterior margin.
Prosternum less than half the length of forecoxae, anterior margin of proster-
num straight. Length of propleura markedly less than their width. Forecoxae
large, their length not less than length of hind coxae. Mesosternum with a
depression on anterior comer and two longitudinal lines in front of middle

45 coxae. Length of metasternum one-fifth its width. Distance between middle
and hind coxae half the length of the former. Distance between lateral corners
of coxae nearly two-thirds the distance between apex of middle coxa and
lateral corner of hind coxa. Abdomen same length as meso- and metathorax
together, almost all its sternites equal in length, only the last slightly longer.

Dimensions. Body length 3.7 mm, width 2.2 mm, elytral length 2.8 mm.

*Probably the eyes are not literally ‘overlapping’, rather the outlines of the dorsal and ventral
eyes are superimposed as in Figure 19—Scientific Editor.

3)

Genus Cretotortor Ponomarenko, 1973

Cretotortor archarensis Ponomarenko, sp. nov.
(Plate III, Photo 8, Figure 20)

Species name coined from Arkhara site.

Holotype. No. 2055/75. PIN, impression of left elytron. Amur oblast,
quarry at Arkhara station (Arkhara site). Upper Cretaceous, Tsagayan.

Material. Holotype and paired elytra of specimen No. 2055/19 from the
same site.

Description. Length of elytron approximately twice the width. Lateral
border wider than intervals between elytral grooves, sutural angle obtuse,
section of lateral comer only a little longer than that of apical. Elytral interval
with transverse rugae and distinct punctures. Suture margined.

Dimensions. Elytral length 6.5—7.1 mm, width 3.2 mm.

Comparison. Distinguished from the second* species of the genus by
the wider lateral margin of elytra, the almost equal sections of its apical
corners and by the distinct punctures on the elytral intervals.

Fig. 20. Cretotortor archarensis, sp. nov.; holotype PIN No. 2055/75. Arkhara,
Upper Cretaceous.

*Not described in this treatment—General Editor.

46

60

INFRAORDER GEADEPHAGA
SUPERFAMILY CARABOIDEA LATREILLE, 1825
Family TRACHYPACHEIDAE Leconte, 1861
Subfamily Eodromeinae Ponomarenko, Subfam. Nov.

Diagnosis. Clypeus not extending to point of antennal attachment. Fore-
coxal cavities open. Lateral wall of middle coxal cavities formed by meso-
sternum, mesepimeron and metasternum. Mesepisternum not extending to
middle coxal cavities, minimum separation equal to mesepisternal mesal
margin. Hind coxae separating metasternum and abdomen, usually with large
femoral plates. Both spurs of foretibiae apical.

Composition. Seven genera in the Triassic, Jurassic and Early Creta-
ceous of Asia, and possibly the Late Jurassic of Western Europe.

Comparison. Distinguished from the nominal subfamily by the
metepisternum extending to middle coxal cavities, while the mesepisternum
are farther removed from them.

Genus Sogdodromeus Ponomarenko, gen. nov.

Genus name coined from the region Sogdiana and ‘dromeus’ (Greek) —
runner.

Type species. S. altus, sp. nov. Triassic of Soviet Central Asia.

Description. Small, flat beetle. Head transverse, length of head capsule
approximately half its maximum width at eyes. Gular plate longer than wide.
Antennae rather long and thick, weakly moniliform. Pronotum strongly
transverse, widest in anterior third, expanded in front of posterior angles.
Mesosternum shorter than the transverse middle coxae. Distance between
middle and hind coxae equal to length of middle coxae. Transverse metaster-
nal suture slightly convex posteriorly. Hind coxae slightly oblique, their
femoral plates large, in medial half only slightly shorter than coxal width,
from this point sharply tapered laterally, but extending to lateral coxal corners
as a fairly wide band. Coxae concealed under femoral plates. Length of
abdomen half that of meso- and metasternum together, its apex pointed;
width of the last sternite half the width of abdominal base. Sternites articulate,
movable, telescoping.

Species Composition. Monotypic genus.

Comparison. Most closely resembles Platycoxa in the possession of
large femoral plates fully covering the coxae. Differs from it by the transverse
middle coxae and shape of laterally sharply tapered femoral plates.

47

61

Sogdodromeus altus Ponomarenko, sp. nov.
(Plate IV, Photo 1; Figure 21)

Species name coined from ‘altus’ (Latin)—ancient.

Holotype. No. 2971/417, PIN, impression of a beetle lacking most parts
of legs and antennae. Soviet Central Asia, Kirgiz SSR, Osh oblast, Batken
region, Madygen area (Madygen site, south-western part). Triassic, Madygen
series.

Material. Holotype.

Description. Head including mandibles much shorter than broad. Man-
dibles short. Eyes shorter than temples, longer than very short genae. Length
of pronotum two-fifths its width, anteriorly roundly emarginate; anterior
angles acute, posterior almost right-angled. Prosternum much shorter than
pronotum, longer than forecoxae, anteriorly with flat, longitudinal extension
and anterior prolongation of the intercoxal process. [Intercoxal] process flat,
broad, broadening behind coxae, truncate apically. Mesosternum slightly
shorter than middle coxae. Length of metasternum approximately one-fourth
its width, its anterior margin half of posterior margin. Metepisterna strongly
broadening toward the anterior margin. Length of femoral plates two-thirds
the width, their elongate mesal part not shorter than the apical width; apex
weakly rounded. Coxae under femoral plates strongly prolonged along
midline of body. Abdomen tapering from the base. First, fifth and sixth
visible sternites equal in length; second and third slightly longer; last one-
and-a-half times longer than the penultimate, its length two-fifths its width
almost triangular. Legs short and weak. Middle femora fusiform, thickened,
widest at middle. Tibiae slender, linear, scarcely broadening apically. Middle
tarsal segments short, apically broadening a little. Body dorsally with small
dense punctation, punctures on elytra fused into transverse rugae. Ventrally
with larger punctures, punctures on meso- and metasternum markedly larger,
abdominal sternites with dense, longitudinal, slightly convoluted striae.

Dimensions. Body length9.5 mm, width 4.5 mm; elytral length 6.5 mm.

Genus Platycoxa Ponomarenko, gen. nov.

Genus name coined from ‘platys’ (Greek)—flat and ‘coxa’ (Greek)—thigh.
Type species. P.armata, sp.nov. Lower Jurassic of Soviet Central Asia.
Description. Small, flat beetles. Head transverse, length of head capsule

less than half its width. Temples and genae shorter than eyes. Antennae rather

long and thick, their segments only slightly longer than wide. Pronotum
strongly transverse, anteriorly not much narrowing. Mesosternum not longer
than middle coxae, distance between middle and hind coxae much greater
than [length of] coxae themselves. Transverse metasternal suture almost
straight. Hind coxae slightly oblique, their femoral plates large, transverse.

62

Fig. 21. Sogdodromeus altus, sp.nov.; holotype PIN No. 2971/417: a—dorsal view;
b—ventral view. Madygen, Triassic.

laterally reduced but extending up to the sides of coxae, here the length of
femoral plates not less than that of the coxae, coxae not projecting from under
femoral plates. Abdomen rather short, less than twice as long as the meso-
and metathorax together, apically pointed, width of the last sternite much less
than that of the base of abdomen.

Species composition. Two species in the Early Jurassic of Soviet
Central Asia and East Kazakhstan.

Comparison. Distinguished from all other species by shape of large
femoral plates of hind coxae, which are laterally not much tapered and extend
to the sides of coxae.

Platycoxa armata Ponomarenko, sp. nov.
(Plate IV, Photo 2; Figure 22a)

Species name coined from ‘armata’ (Latin)—armed.

Holotype. No. 2903/222, PIN, impression of a beetle lacking antennae
and legs. Kirgiz SSR, Ton region, south bank of Issyk-Kul’ meander* to the
south of Kadzhi-Sai settlement (Issyk-Kul’ site). Lower Jurassic, Dzhil’
series.

Material. Holotype and impression of metathorax and abdomen of a
beetle PIN No. 371/498 from the same site.

*In the Russian original abbreviated as Iz. for ‘izvilina-—General Editor.

49

63

Description. Length of head with mandibles greater than length of
prothorax, two-thirds the width of occiput. Length of the head capsule
two-fifths its width, sharply narrowing anteriorly in front of eyes. Eyes large,
almost twice as long as the temples, genae very short. Mandibles long,
upright, longer than the head capsule, sharply curved apically. Pronotum
almost rectangular, slightly narrower than elytra at shoulders, anteriorly
weakly emarginate, its length less than half the width. Posternum in front of
forecoxae shorter than coxal length; prosternal process noticeably extending
beyond coxae, half the coxal width, apex rounded. Mesosternum shorter than
middle coxae. Length of metasternum two-fifths its width at posterior mar-
gin, its anterior margin almost one-third the posterior; posterior margin very
slightly angularly projecting backward; transverse metasternal suture

Fig. 22. Representatives of genus Platycoxa. a—P. armata sp. nov., holotype PIN.
No. 2903/222, Issyk-Kul’, Lower Jurassic; b—P. jurassica sp. nov., holotype PIN
No. 2496/6, Kenderlyk, Lower Jurassic.

49

64

straight. Distance between middle and hind coxae twice the length of middle
coxae. Width of femoral plates of hind coxae 1.7 times the length, tapered in
lateral half; their length in lateral third approximately half the maximum
length. Abdomen one-and-a-half times longer than meso- and metathorax
together, narrowing from the base of fourth sternite; width of last sternite 2.7
times its length, half the width of abdominal base. All abdominal sternites
subequal in length. Body with coarse, dense, fused punctation; punctures
particularly large on metasternum where they form transverse rugae.

Dimensions. Body length 7.1 mm, width 2.8-3.0 mm; elytral length
5.0-6.0 mm.

Comparison. Distinguished by long crescent-shaped mandibles; long,
coarsely punctate metasternum; and less oblique hind coxae.

Platycoxa jurassica Ponomarenko, sp. nov.
(Plate IV, Photo 3; Figure 22b)

Holotype. No.2496/6, PIN, impression of beetle without apices of antennae,
middle legs, fore- and hind tarsi. East-Kazakhstan oblast, Zaisan region, Saur
range, right bank of Akkolka river, tributary of Karaungir river (Kenderlyk
site), Lower Jurassic, Tologoi series.

Material. Holotype.

Description. Length of head half the width of occiput, sharply narrow-
ing in front of eyes. Genae and temples very short. Gular plate twice as long
as wide, much narrower than submentum; lateral lobes of submentum short,
round, larger than median lobe. First antennal segment noticeably longer than
third which is not longer than fourth; second segment transverse. Pronotum
twice as long as wide, anteriorly roundly emarginate, anterior and posterior
angles acute, drawn out [away from midline]; lateral margin posteriorly
weakly emarginate at middle. Prosternum longer than small forecoxae;
prosternal process less than half the width of coxae, distinctly not extending
to their posterior margin. Propleura almost triangular, slightly longer than
wide. Mesosternum not shorter than middle coxae, in front with triangular
depression for prosternal process. Mesepisterna almost rectangular, mese-
pimera short, taenioid. Length of metasternum one-third its width at posterior
margin, its anterior margin half the posterior. Distance between middle and
hind coxae one-and-a-half times the length of middle coxae, posterior
metasternal margin angularly projecting backward. Metepisterna strongly
broadening in anterior third, width of its anterior margin half the length.
Length of femoral plates of hind coxae half the width, longest in the mesal
third and gradually tapering laterally from there; lateral length of plates
one-third their maximum length; posterior coxal margin scarcely peeking
from beneath plates at lateral margins. Forefemora noticeably shorter than
hind femora, slightly thickened uniformly. Foretibiae slightly shorter than

50

65

femora, curved, slightly broadened at apex. Hind femora widest in the
proximal third, weakly drawn out before apex, three times as long as
trochanters. Hind tibiae slender, almost linear, one-fourth longer than
femora. Body with small punctures; punctures noticeably larger on metaster-
num and metepisterna.

Dimensions. Body length 5.0 mm, width 2.5 mm; elytral length. 7mm.

Comparison. Distinguished by shorter mandibles, shorter metaster-
num, and laterally strongly tapering femoral plates.

Genus Unda Ponomarenko, gen. nov.

Genus name coined from Unda River.

Type species. U. microplata, sp. nov. Lower Cretaceous of Trans-
Baikal.

Description. Small beetles. Head transverse, triangular. Eyes small, not
longer than temples. Antennae rather long, slender, filiform. Pronotum
transverse, with weak emargination in front, slightly tapering in anterior
third. Elytra at shoulders broader than pronotum. Mesosternum nearly equal
to middle coxae in length, distance between middle and hind coxae much
greater than [length of] middle coxae. Transverse, metasternal suture an-
gularly projecting backward. Hind coxae rather strongly oblique; their
femoral plates transverse laterally tapering, extending as narrow tongue up
to lateral third of coxa which is posteriorly visible from under them. Ab-
domen rather short, only slightly longer than meso- and metathorax together,
apex pointed; last sternite longer than the remaining, its anterior margin much
narrower than base of abdomen.

Species composition. Three species in the Early Cretaceous of Trans-
Baikal.

Comparison. Differs in the shape of the femoral plates of hind coxae,
which are much shorter than wide, gradually tapering laterally, without
notch, and extending up to lateral third of coxae.

Unda microplata Ponomarenko, sp. nov.
(Plate IV, Photo 4; Figure 23)

Species name coined from ‘micros’ (Greek)—small; and ‘platys’ (Greek) —
flat.

Holotype. No. 3015/371, PIN, impression of beetle lacking most parts
of legs. Trans-Baikal, Chita oblast, Balei region, right bank of Unda river, 2
km upstream of Zhidka settlement (Unda site). Lower Cretaceous, Aptian-
Albian?, Balei series.

Material. Holotype.

5)

66

Description. Head together with mandibles slightly shorter than width
of occiput, width of occiput 1.8 times the length of head capsule, the latter
narrowing anteriorly from base. Eyes same length as temples, twice as long
as genae. Length of gular plate twice its width. Antennae extending beyond
the base of pronotum by five segments, segments slightly broadened apically,
three apical segments thicker than the rest, distinctly moniliform. First
antennal segment equal in length to the third; the second equal to half, and
the fourth to two-thirds the third; fourth and fifth longer than sixth to eighth;
ninth to eleventh equal in length to the fourth; the last segment teardrop-
shaped, pointed apically, its length twice the width. Mandibles short, apically
curved. Width of pronotum 1.7 times the length, anteriorly very weakly
roundly emarginate. Prosternum 1.5 times longer than forecoxae; prosternal
process short, noticeably narrowing posteriorly. Mesosternum with distinct
triangular depression for prosternal process, mesepisterna almost rectan-
gular, transverse; mesepimera short, slightly laterally broadened. Length of

Fig. 23. Unda microplata, sp. nov,; holotype PIN No. 3015/371. Unda, Lower
Cretaceous.

3}

—

67.

metasternum approximately one-third its width at posterior margin, anterior
margin narrower than posterior, posterior margin strongly angularly project-
ing backward, transverse metasternal suture angular. Distance between mid-
dle and hind coxae one-and-a-half times greater than [length of] the middle
coxae. Mesepisternum strongly broadening in anterior third, its length 1.7
times the width at anterior margin. Hind coxae short, slightly tapering
laterally, femoral plates only slightly longer than coxae, not extending to
posterior margin of the second abdominal segment, longest in the mesal third,
from where it gradually tapers laterally; in the lateral third extending as a
very narrow tongue almost up to lateral coxal margins. First abdominal
sternite two-thirds the length of second, second longer than third and only
slightly shorter than the terminal. Abdomen slightly broadening toward
fourth sternite, and then narrowing posteriorly. Base of last sternite two-
thirds the base of fourth, twice the length of the last sternite. Sternal lobes of
the true eighth segment triangular, valves extended, apically pointed with
small stylus. Hind trochanters very broad; hind femora clavate, dilated in
apical third; segments of middle and hind tarsi slightly broadening toward
apex, not very long. Body with rather dense, small punctures.

Dimensions. Body length 3.8 mm, width 1.9 mm; elytral length 3.0 mm.

Comparison. Distinguished by the long prosternum and shape of
femoral plates which are laterally gradually tapered.and lacking a distinct
postero-lateral angle.

Unda angulata Ponomarenko, sp. nov.
(Plate IV, Photo 5; Figure 24)

Species name coined from ‘anguiata’ (Latin)—angular.

Holotype. No. 2372/24, PIN, impression of almost entire beetle but
dismembered into individual sclerites. Chita oblast, Shelopugin region, left
bank of Daya river above Shiviya ravine (Daya site). Lower Cretaceous,
Aptian-Albian. Balei series.

Material. Holotype.

Description. Length of head slightly shorter than width of occiput,
length of gular plate twice its width. Antennae not shorter than head and
pronotum together, their segments short, distinctly moniliform, last segment
oval, apically rounded. Prosternum in front of forecoxae only a little longer
than them; propleura rather wide. Mesosternum shorter than middle coxae,
mesepisternum almost triangular, mesepimeron markedly broadening at
lateral margin. Length of metasternum two-fifths its width at posterior
margin; posterior margin strongly angularly projecting backward, transverse
metasternal suture less strongly projecting. Distance between middle and
hind coxae almost twice the length of middle coxae. Metepisterna gradually
broadening in front of middle, their width at anterior margin two-fifths the

68

Fig. 24. Unda angulata, sp. nov.; holotype PIN No. 2372/24. Daya, Lower
Cretaceous.

length. Hind coxae short, slightly tapering laterally. Femoral plates not
longer than coxae, their length two-thirds the width, posterior margin in
mesal two-thirds truncate at right-angle to body axis; more laterally the
femoral plates sharply tapered, extending up to the lateral fourth; postero-
lateral angle distinct, almost straight. Abdomen approximately equal in
length to meso- and metathorax together, widest at the fourth sternite; basal
width of the last sternite half of abdominal base, almost three times wider
than long. Foretrochanters equal in length to coxae, one-third the length of
forefemora. Forefemora rather thick, widest in apical third, distally narrow-
ing. Tibia a little shorter than femur, somewhat broadening apically; apex of

52 tibia broad and notched, forming an organ for cleaning antennae. Spurs of

foretibiae a little longer than first tarsal segment. Tibiae with a row of rather
long, stiff spines. Tarsus two-thirds the length of tibia, its first segment
thicker and longer than the second, second a little longer than the third and
fourth, the last equal to the two preceding together. Middle and hind femora
equal in length to forefemora, uniformly broadened, width four times the
length. Tibiae slender, linear, one-fourth longer than femora; spurs of middle
tibiae slender and short, shorter than first tarsal segment. Middle tarsi
two-thirds the length of tibiae, hind tarsi almost equal to them. Tarsal
segments, slender, linear. First segment of middle tarsi equal to the second
and third together, second longer than third, fourth equal to second, fifth
slightly shorter than first. Elytra smooth, body without distinct punctation.

53

69

Dimensions. Body length about 6.0 mm, width 2.5 mm; elytral length
4.5 mm.

Comparison. Distinguished by femoral plates with a distinct angle at the
postericr margin.

Unda cursoria Ponomarenko, sp. nov.
(Plate IV, Photo 6; Figure 25)

Species name coined from ‘cursor’ (Latin)—runner.

Holotype. No. 2372/23 PIN, impression of an entire beetle, distal seg-
ments of fore- and middle legs bent under body and not visible. Trans-Baikal,
Chita oblast, Shelopugin region, left bank of Daya river above Shiviya ravine
(Daya site). Lower Cretaceous, Aptian-Albian, Balei series.

Material. Holotype.

Description. Head including mandibles shorter than wide, head capsule
almost rectangular, narrowing only in front of eyes, temples a little longer
than eyes, genae very short, antennae inserted at anierior margin of eyes.
Gular plate longer than wide, anteriorly broadening. Only apical antennal
segment extending beyond pronotal base. First segment dilated, twice as long
and one-and-a-half times as thick as doliform second; third and fourth equal

Fig. 25. Unda cursoria, sp. nov.; holotype PIN No. 2372/23; a—dorsal view;
b—ventral view; c—head; d—metasternum. Daya, Lower Cretaceous.

35)

70

to each other and three-fourth the length of first; fifth slightly shorter; sixth
to eighth markedly shorter than remaining, almost equal in length and width,
last ovoid. Labrum weakly notched apically. Mandibles small, upright,
retinaculum weak. Submentum strongly broadening anteriorly; lobes of
mentum small, rather narrow, lateral ones larger than middle. Maxillary palps
much longer than mandibles. Pronotum almost rectangular, slightly broaden-
ing toward anterior third and then slightly narrowing beyond, anterior margin
very weakly roundly emarginate, anterior angles rounded, posterior margin
slightly projecting backward. Disk of pronotum raised, with median long-
itudinal groove. Prosternum very short, almost half the forecoxae; prosternal
process narrow, extending a little beyond forecoxae, apically rounded.
Propleura narrow, twice as iong as wide, jateral margin with emargination
behind middle; beyond coxae propleuron forming mesally directed short
process. Mesosternum short, transverse, half the length of middle coxae,
without distinct depression for prosternal process. Mesepisterna almost equal
in length and width, rectangular; mesepimera narrow, laterally almost not
broadening, taenioid. Middle coxae convergent. Length of metasternum
one-third its width at the posterior margin, its anterior margin half of the
posterior; distance between middle and hind coxae a little greater than length
of middle coxae. Metepisternum strongly and rather sharply broadening in
anterior part, its width at anterior margin two-and-a-haif times the medial
width. Hind coxae strongly reduced in length beyond mesal fourth. Femoral
plates not longer than hind coxae, very slightly less in length than width,
laterally extending slightly beyond coxal middle, their lateral margins form-
ing an almost straight line parallel to body axis; antero-lateral process absent,
postero-lateral angle rounded. Abdomen a little longer than meso- and
metathorax together, broadening from base to apex of second visible sternite,
then narrowing; third to fifth sternites narrow; width of the last 3.5 times its
length, its base two-thirds the abdominal base. Elytra smooth, lateral border
narrow. Legs long, femoral apices rather strongly extending beyond lateral
body margins. Femora very slightly thickened, forefemora shorter than
subequal middle and hind femora. Hind tibiae almost one-fifth longer than
femora slender, almost not broadening apically. Hind tarsi four-fifths the
tibiae, their first and last segments longer than middle segments, slender,
linear. Body smooth, punctation indistinct.

Dimensions. Body length 7.0 mm, width 3.5 mm; elytral length 4.5
mm.

Comparison. Close to U. angulata in many characters, but distin-
guished by the short prosternum, anteriorly strongly broadening metepisterna
and narrower femoral plates of hind coxae with rounded postero-lateral
angles.

a A

54

71
Genus Psacodromeus Ponomarenko, gen. nov.

Genus name coined from ‘psakas’ (Greek)—drop, and “dromeus’ (Greek) —
runner.

Type species. P. gutta, sp. nov. Upper Jurassic of South Kazakhstan.

Description. Medium- and small-sized beetles with teardrop-shaped
body. Head including mandibies approximately equal in length and width,
triangular, narrowing anteriorly from base. Temples almost as long as eyes,
genae much shorter. Gular piate much longer than wide. Antennae long and
thick, extending far beyond pronotal base. Pronotum strongly roundly taper-
ing anteriorly from base, at base scarcely narrower than elytra at shoulders.
Mesosternum not longer than middle coxae, distance between middle and
hind coxae only a little more than the length of middle coxae. Hind coxae
with large femoral plates, length of which not less than width; posterior
margin of plates straight or rounded and without emargination, lateral emar-
gination shallow. Abdomen less than twice the length of meso- and
metathorax together, pointed apically; width of last sternite much less than
width of abdominal base. Elytra smooth.

Species composition. Four species in the Late Jurassic of South
Kazakhstan.

Comparison. Distinguished from other genera possessing long femoral
plates of the hind coxae by the streamlined teardrop-shaped body; laterally,
from elytra to head, the body practically describes a single smooth curve.

Psacodromeus gutta Ponomarenko, sp. nov.
(Plate IV, Photo 7; Figure 26)

Species name coined from ‘gutta’ (Latin)—drop.

Holotype. No. 2554/456, PIN, almost entire impression of male. South
Kazakhstan, Chimkent oblast, Algabass region south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Besides the holotype, two aimost entire impressions of beet-
les, specimen Nos. 2066/2514 and 2066/2865 from the same site, and the
body of a beetle without head and prothorax, specimen No. 2542/279 from
the Karatau-Galkino site.

Description. Head capsule almost half the width of occiput, temples
slightly shorter than eyes. Anterior margin of labrum straight. Length of gular
plate twice the width. Antennae extending beyond pronotal base by four to
five segments. First segment a little thicker than the remaining segments, in
length approximately equal to the third, second approximately half that
length, fourth noticeably shorter than third, fifth to ninth equal and shorter
than fourth, tenth not longer than second, last ovoid. Length of pronotum half

a | fh

\ ae,
\ Za \

: \

\

54 Fig. 26. Psacodromeus gutta, sp. nov.; holotype PIN No. 2554/456: a—dorsal
view; b—ventral view. Karatau, Upper Jurassic.

the basal width, anterior margin two-thirds the width of posterior. Anterior
margin of pronotum with shallow rounded emargination. Prosternum
noticeably shorter than pronotum; prosternal length in front of forecoxae
noticeably less than coxal length. Prosternal process apically rounded, its
width half that of forecoxae. Mesosternum slightly shorter than middle
coxae, with small triangular depression for prosternal process. Mesopleura
markedly shifted anteriorly in relation to mesosternum; mesepisterna almost
rectangular; mesepimera rather long, slightly broadening laterally. Length of
metasternum two-fifths the width at posterior margin, width at posterior
margin 1.7 times the anterior. Distance between middle and hind coxae
nearly equal to length of the middle coxae. Posterior margin of metasternum
slightly angularly projecting backward, transverse metasternal suture almost
straight. Metepisterna strongly but not sharply broadening in anterior third.
Hind coxae laterally sharply tapering in mesal fourth, nearly untapered

73

55 beyond; mesal margin almost twice the length of lateral margin. Femoral
plates more than twice the length of hind coxae, extending up to mid-
abdomen, almost twice as long as medial width; basally drawn out into
narrow tongue in the lateral half of coxa and extending to lateral corners of
metasternum. Posterior margin of femoral plate rounded, laterally with a
shallow emargination. Abdomen 1.7 times longer than meso- and metaster-
num together, narrowing from the commencement [anterior margin] of fourth
sternite; base of last sternite half the width of abdominal base, three times its
own length. Legs rather long, femora extending by one-third beyond lateral
body margins, gradually broadening toward apical third, distally narrowing,
slightly drawn out before apex; forefemora two-thirds the length of middle
femora and five-eighths the hind femora. Tibiae very slightly broadening in
distal third, foretibiae equal in length to femora, middle and hind tibiae
slightly shorter than the corresponding femora, apical spurs of foretibiae
small; groove for cleaning antennae oblique. Foretarsi equal in length to
tibiae, their basal segments almost equal, apical segment longer than them.
Middle tarsi scarcely longer and hind tarsi noticeably longer than the cor-
responding tibiae; their first segments longer than spurs of tibiae; segments
of middle tarsi equa! in length; in hind tarsi last segment longer than the
remaining [= preceding] segments. Body with small punctures.

Dimensions. Body length 9.8-10.8 mm, width 4.5-5.3 mm; elytral
length 6.8—7.7 mm.
Comparison. Distinguished by long femoral plates of hind coxae.

Psacodromeus ovalis Ponomarenko, sp. nov.
(Plate IV, Photo 8; Figure 27)

Species name coined from ‘ovalis’ (Latin)—oval.

Holotype. No. 2997/1849, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Besides holotype, almost entire impression of beetle,
specimen No. 2384/757 from the same site.

Description. Length of head capsule two-thirds the width. Temples
scarcely shorter than eyes. Anterior margin of labrum slightly emarginate.
Length of gular plate two-and-a-half times the width. Antennae extending
beyond pronotal base by four segments. First segment thicker and two-thirds
the third, second half the length of third, fifth slightly longer than first, fourth
equal to first, remaining except the tenth equal to fifth, tenth slightly shorter,
last ovoid. Pronotum anteriorly with rather deep straight emargination, its
anterior margin five-sevenths the posterior, length half the basal width.
Prosternum slightly shorter than pronotum; length of prosternum in front of

74

5 —— fb

Fig. 27. Psacodromeus ovalis, sp. nov.; holotype PIN No. 2997/1849: a—dorsal
view; b—ventral view. Karatau, Upper Jurassic.

forecoxae very slightly more than coxal length. Prosternal process straight,
apically rounded, scarcely longer than hind coxae, two-thirds their width.
Mesosternum half the length of middle coxae, triangular depression for
prosternal process rather large. Mesopleura somewhat shifted anteriorly in
relation to mesosternum, mesepisterna almost rectangular, mesepimera
rather long, laterally slightly broadening. Length of metasternum almost
one-third its width at posterior margin, width at posterior margin 1.7 times
at anterior; distance between middle and hind coxae slightly more than the
length of middle coxae. Posterior margin of metasternum very slightly
angularly projecting backward; transverse metasternal suture almost straight.
Metepisterna sharply broadening in front of anterior margin. Length of hind
coxae laterally strongly reduced from mesal fourth, less than half at the lateral
than at the mesal margin. Femoral plates 1.7 times longer than coxae,
abruptly shortened at midwidth and extending to comers of metasternum as
a narrow angular process bordering the anterior coxal margin. Posterior
56 margin of the elongate part of coxa almost straight, postero-lateral angle well
defined, shape of postero-lateral part almost rectangular. Abdomen one-and-
a-half times longer than meso- and metasternum together, narrowing from

Te)

base of second sternite, basal width of the last sternite half of abdominal base,
length of last sternite one-fourth its width. Not more than one-fourth of
femora extending beyond lateral body margins, femora rather weakly
broadening toward middle, distally narrowing; forefemora very slightly
shorter than middle femora, four-fifths the length of hind femora. Tibiae very
slightly broadening in distal fourth, fore- and middle tibiae two-thirds the
corresponding femora, hind tibiae equal to hind femora. Foretarsi not shorter
than tibiae. Body with small dense punctures.

Dimensions. Body length 7.2-8.3 mm, width 3.8-4.0 mm; elytral
length 5.2-5.7 mm.

Comparison. Distinguished by the shorter and almost rectangular
femoral plates.

Psacodromeus crassus Ponomarenko, sp. nov.
(Plate V, Photo 1; Figure 28)

Species name coined from ‘crassus’ (Latin)—thick.

Holotype. No. 2066/3147, PIN, impression of a beetle without antennae
and legs. South Kazakhstan, Chimkent oblast, Algabass region, south-
western border of Kashkarata river valley, Aulie area near Mikhailovka
village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

ane

Hons
i
We
ee, 3

Fig. 28. Psacodromeus crassus, sp. nov.; a—holotype PIN No. 2066/3147,
b—paratype PIN No. 2066/2349. Karatau, Upper Jurassic.

Sil

76

Material. Besides holotype, impression of beetles lacking legs and a
large part of antennae, specimen Nos. 2066/2349, 2904/879, 2997/405 from
the same site.

Description. Head including mandibles slightly longer than width of
occiput; length of head capsule slmost equal to width; temples not shorter
than eyes. First antennal segment slightly thicker and very slightly shorter
than third, second half the length of third, third noticeably longer than fourth
and fifth. Pronotum anteriorly with rather deep straight emargination, prono-
tal length half the width; posterior margin 1.3 times the anterior. Length of
prosternum in front of forecoxae noticeably more than coxal diameter.
Prosternal process extending slightly beyond forecoxae and half their width.
Mesosternum half the length of middle coxae, triangular depression for
prosternal process rather large. Mesopleura strongly shifted anteriorly in
relation to mesosternum. Length of metasternum one-third the width at
posterior margin; anterior margin half the width of posterior, distance be-
tween [middle and hind] coxae greater than length of middle coxae. Posterior
margin of metasternum angularly projecting backward, transverse metaster-
nal suture distinctly angular. Metepisterna sharply broadening in anterior
third. Femoral plates of hind coxae strongly shortened in middle. Narrow
tongue-like process of anterior margin not extending to lateral corners of
metasternum. Length of the elongate part of the femoral plate slightly more
than width, its posterior margin rounded. Abdomen slightly longer than
meso- arid metasternum together; abdominal base 1.7 times the base of last
sternite, length of last sternite one-third its width. Body with fine, sparse
punctation.

Dimensions. Body length 6.5—7.2 mm, width 2.9-3.4 mm; elytral
length 4.2-4.5 mm.

Comparison. Distinguished by long head and rounded femoral plates.

Psacodromeus rugosus Ponomarenko, sp. nov.
(Plate V, Photo 2; Figure 29)

Species name coined from ‘rugosus’ (Latin)—wrinkled.

Holotype. No. 1789/214, PIN, almost entire impression of a beetle.
South Kazakhstan, Chimkent oblast, Algabass region, Donner-Bulak area
near Uspen settlement (Karatau-Galkino site). Upper Jurassic Karabastau
series.

Material. Holotype.

Description. Length of head including mandibles equal to width of
occiput; head capsule twice as long as mandibles; temples slightly shorter
than small eyes. Length of gular plate twice the medial width. Antennae
extending beyond pronotal base by four segments. First antennal segment
scarcely thicker than the remaining, length of the first segment together with

58

A

second equal to third; third one-and-a-half times longer than fourth. Length
of pronotum five-ninths the width at posterior margin; anterior margin two-
thirds the width of posterior, with deep rounded emargination. Prosternum
small, shorter than forecoxae, with flat keel in the middle—the continuation
of the prosternal process; propleura large, their length one-and-a-half times
the width. Forecoxae rounded, contiguous under the narrow prosternal
process. Prosternal process rounded apically, two-fifths the width of coxae,
its apex scarcely extending beyond coxae. Mesosternum narrow, mesopleura
strongly shifted anteriorly. Length of metasternum one-third the width at
posterior margin, anterior margin half the posterior; posterior margin strong-
ly angularly projecting backward; distance between middle and hind coxae
almost one-and-a-half times more than the length of middle coxae.
Metepisternum in anterior half strongly but not sharply broadening. Hind
coxae shortened to about half at mesal fourth, further laterally almost equal
in length; anterior coxal margin convex, strongly curved backward toward

)
=
8

LD Bese
als

ne

Fig. 29. Psacodromeus rugosus, sp. nov.; holotype PIN No. 1789/214: a—ventral
view; b—dorsal view. Karatau, Upper Jurassic.

59

78

middle of body. Length of femoral plates sharply reduced laterally from the
middle of coxae, their narrow tongue-like lateral process basally short. not
extending to corners of metasternum. Posterior margin of femoral plates
rounded, laterally without emargination. Abdomen only slightly longer than
meso- and metasternum together, narrowing from the anterior margin of
fourth sternite. Legs rather long, femoral apices noticeably extending beyond
lateral body margins. Femora thickened, broadening in distal half, forefemur
three-fourths the middle femur, two-thirds the hind femur. Foretibia slightly
shorter than femur, noticeably broadening in distal half, notch extending to
middle of tibia, spurs of foretibia longer than first tarsal segment. Three basal
segments of foretarsus broadened, almost one-and-a-half times longer than
wide. Tarsal length apparently almost equal to tibia. Middle tibiae slightly
shorter than femur, equal to tarsus; tarsal segments slender, almost equal.
Length of hind tibiae equal to femur, slender, slightly broadening from base
to apex; tarsus slightly shorter than tibia, its segments slender, linear. Body
with rather large punctures, particularly coarse on elytra, pronotum and
metasternum.

Dimensions. Body length 10.7 mm, width 5.4 mm; elytral length 6.8
mm.

Comparison. Occupies isolated position among other species of the
genus, differs in anteriorly deeply emarginate pronotum, small prosternum
with keel, narrow prosternal process, strongly oblique hind coxae, longer legs
and distinct punctures.

Genus Karatoma Ponomarenko, gen. nov.

Genus name coined from Karatau mountain and ‘tome’ (Greek)—cut.

Type species. K. agilis, sp. nov. Upper Jurassic of South Kazakhstan.

Description. Rather large, flat beetles with long legs and mandibles.
Length of head including mandibles much longer than width of head capsule;
the latter equal to width of head near base, narrowing anterior to eyes; length
of temples not less than length of eyes. Antennae long and slender. Pronotum
wide, not narrower than elytra at shoulders, very weakly narrowing in
anterior third, anteriorly weakly emarginate, anterior angles acute. Metaster-
num short; distance between middle and hind coxae not longer than middle
coxae. Hind coxae with large femoral plates, posterior margin of femoral
plates crenate, lateral margin with backward projection straight. Abdomen
slightly longer than meso- and metasternum together; basal width of the last
sternite two-thirds the abdominal base; apex of abdomen rounded, with small
emargination. Elytra with large irregularly distributed punctures. Legs long,
hind tibiae longer than femora.

Species composition. One species in the Upper Jurassic of South
Kazakhstan and one species in the Lower Cretaceous of Trans-Baikal.

60

79

Comparison. Distinguished from other genera with similar shape of
femoral plates by the broad, almost rectangular pronotum, long legs, and
coarse punctures.

Karatoma agilis Ponomarenko, sp. nov.
(Plate V, Photo 3; Figure 30)

Species name coined from ‘agilis’ (Latin)—quick.

Holotype. No. 2784/1528, PIN, impression of a beetle without distal
segments of forelegs. South Kazakhstan, Chimkent oblast, Algabass region,
south-western flank of Kashkarata river valley, Aulie area near Mikhailovka
village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype. An incomplete impression of a beetle, specimen
No. 2997/2442 was found in the same site; of which only the elytra were
intact from under which the apices of very long femora and bases of tibiae
were visible. Judging from the characteristic features (very coarse punctures,
depigmented spots in the apical part of elytra, long legs), the fragment may
belong to a beetle of the same species, but its dimensions are almost one-third
larger.

Description. Length of head including mandibles one-and-a-half times
greater than length of prothorax or width of occiput; temples and eyes equal

b

Fig. 30. Karatoma agilis, sp. nov.; holotype PIN No. 2784/1528: a—dorsal
view; b—ventral view. Karatau, Upper Jurassic.

60

80

in length, genae half of them. Antennae extending beyond pronotal base by
three segments. First antennal segmenta little thicker than flagellar segments,
together with second segment equal in length to third; fourth very slightly
longer than fifth and noticeably shorter than third. Flagellar segments almost
cylindrical, only slightly broadening at apices. Labrum weakly emarginate
at apex. Mandibles long, crescent-shaped, asymmetrical, retinaculum small.
Width of pronotal base 1.7 times the pronotal length; anterior pronotal angles
extending forward; disk of pronotum with raised transverse, oval portion
divided by a median longitudinal groove; sides of pronotum not margined.
Length of metasternum one-fourth its width at posterior margin, width of
posterior margin 1.7 times the anterior; distance between [middle and hind]
coxae equal to length of middle coxae, posterior margin of metasternum
slightly angularly projecting backward. Transverse metasternal suture almost
straight. Femoral plates noticeably longer than coxae, sharply reduced in
length at middle; their antero-lateral tongue-like process long, extending up
to corners of metasternum. Last abdominal sternite longer than the remain-
ing, two preceding sternites shortest. Femora uniformly thickened almost
along the entire length, forefemora slightly shorter than middle femora,
middle femora shorter than hind. Middle and hind tibiae very slender, rather
sharply broadening only at apex, middle tibiae shorter than corresponding
femora, hind tibiae markedly longer. Spurs of middle and hind tibiae shorter
than basal tarsal segment. First segment of middle tarsus scarcely broadening
toward apex, three times longer than the second. Integument with coarse and
dense punctation, elytra with large punctures, medial and apical depigmented
spots on lateral margins of elytra without distinct sculpture*, bottom
[= underside?] with finer punctation.

Dimensions. Body length 19 mm, width 8 mm; elytral length 12 mm.

Comparison. Distinguished by sharply extended anterior angles of
pronotum. Weakly developed raised portion [on disk of pronotum],** long
femoral plates, and depigmented spots on the lateral margins of elytra.

Karatoma raptor Ponomarenko, sp. nov.
(Plate V, Photo 4; Figure 31)

Species name coined from ‘raptor’ (Latin)—robber.

Holotype. No. 3015/362, PIN, impression of a beetle without apices of
antennae and parts of legs. Trans-Baikal, Chita oblast, Balei region, right
bank of Unda river 2 km upstream of Zhidka settlement (Unda site). Lower
Cretaceous, Aptian-Albian?, Balei series.

*Given as structure’ in the Russian original—General Editor.

**Given as ‘weakly expressed relief’ in the Russian original; our interpretation seems
reasonable based on the relevant figures and the complete descriptions of the two Karatoma
spp.—Scientific Editor.

6

6

1

81

Material. Holotype.

Description. Head including mandibles one-third longer than width of
occiput, head capsule slightly shorter than occiput; temples slightly longer
than eyes and genae noticeably shorter than them. First antennal segment
slightly thicker than flagellar segments, first segment together with second
equal in length to the third, second one-third the length of the third; fourth
and fifth segments two-thirds the third. Flagellar segments very slightly
broadening from base to apex of segment. Mandibles long, slightly incurved,
their length almost three times the width; retinaculum small, asymmetric.
Width of pronotum 1.7 times its length, anteriorly with trapezoidal emargina-
tion, anterior angles almost right-angled, disk of pronotum with transverse
raised portion, distant from its posterior margin; with median longitudinal
groove anteriorly joining into triangular depression. Sides of pronotum not
margined. Mesosternum rather long, only slightly shorter than middle coxae,
triangular depression for prosternal process indistinct. Propleura large, al-
most rectangular, pleural suture oblique, mesepisterna and mesepimera al-
most equal. Middle coxae transverse, teardrop-shaped, convergent. Length
of metasternum two-sevenths the width at posterior margin; width of
posterior margin 1.7 times the anterior. Distance between middle and hind
coxae equal to length of the middle coxae. Posterior margin of metasternum
slightly angularly projecting backward, transverse metasternal suture almost
straight. Length of hind coxae strongly reduced laterally, length medially half

_ the maximum length. Femoral plates scarcely longer than hind coxae, almost

N

equal in length and width, occupying somewhat more than half the caxal
width, their lateral margin almost straight; antero-lateral process very weakly
developed, posterior margin with a slight emargination slightly more mesal
to middle, postero-lateral corner rounded. Femora uniformly slightly thick-
ened along the entire length, middle femora slightly broader and shorter than
hind femora. Foretibiae very slightly shorter than femora, slightly broadening
toward apical fourth, grooves for cleaning antennae short. Middle tibiae
scarcely longer than middle femora, weakly curved and broadening from
base to apex. Hind tibiae not broader than middle tibiae, noticeably longer
than hind femora, broadening only in apical fourth. Lateral margins of middle
and hind tibiae with spines; apical spurs shorter than first tarsal segment.
Foretarsi equal to corresponding tibiae, middle and hind tarsi noticeably
shorter than them. Four basal segments of foretarsi short, somewhat broaden-
ing, only slightly longer than wide, last segment narrow, twice as long as the
preceding. Middle tarsi three-fourths the length of tibiae, first segment
slightly broadened, one-and-a-half times longer than the second, third and
fourth equal to second, last segment twice as long as fourth; claws large,
straight, scarcely shorter than the fourth segment. Hind tarsi four-fifths the
tibiae; first and last segments equal, twice as long as the middle segment;
claws large, half the length of middle segment. Integument with coarse

82

punctation. Punctures on head and prosternum smaller and denser, on
metasternum larger and sparser; elytra with large, shallow punctures forming
longitudinal rows transversely interconnected, intervals between them form-
ing convoluted transverse rugae. Anterior margin of pronotum with dense
longitudinal furrows.

Dimensions. Body length 10.7 mm, width 5.2 mm; elytral length 6.7

Fig. 31. Karatoma raptor, sp. nov.; holotype PIN No. 3015/362: a—dorsal view;
b—ventral view; c—head; d—meso- and metathorax, Unda, Jurassic.

83

Comparison. Distinguished by less acute anterior angles of pronotum,
with distinct transverse raised protion on it, and shorter femoral plates of hind
coxae.

Genus Karadromeus Ponomarenko, gen. nov.

Genus name coined from Karatau mountains and ‘dromeus’ (Greek)—run-
ner.

Type species. K.rostratus sp.nov. Upper Jurassic of South Kazakhstan.

Description. Smali, flat beetles. Head including mandibles a little longer
than its basal width, narrowing in front of eyes, length of temples less than
length of eyes. Antennae long, rather thick. Pronotum almost rectangular,
narrowing only in anterior third, anteriorly very weakly roundly emarginate;
its anterior angles acute, extended. Pronotal base much narrower than elytral
base. Metasternum rather large, distance between middle and hind coxae
greater than the length of middle coxae. Hind coxae with large femoral plates,
laterally gradually tapering and extending by narrow tongues almost to the
sides of coxae; medial width of femoral plate jess than its length. Abdomen
a little longer than meso- and metathorax together. Elytra smooth. Legs short,
femora scarcely extending beyond lateral margins of body, tibiae shorter than
femora.

Species composition. Two species in the Upper Jurassic of South
Kazakhstan, and one species each in the Lower Cretaceous of Trans-Baikal
and Mongolia.

Comparison. Distinguished from other genera by the anteriorly slightly
narrowing pronotum, base of which narrower than elytra, and by the shape
of femoral plates of hind coxae.

Karadromeus rostratus Ponomarenko, sp. nov.
(Plate V, Photo 5; Figure 32)

Species name coined from ‘rostratus’ (Latin)—big-nosed.

Holotype. No. 2904/873, PIN, impression of a beetle without distal parts
of antennae and legs. South Kazakhstan, Chimkent oblast, Algabass region,
south-western flank of Kashkarata river valley, Aulie area near Mikhailovka
village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Length of head capsule slightly more than two-thirds its
basal width; eyes one-third longer than temples; genae long, not shorter than
temples. Segments of antennae noticeably broadening from base to apex; first
segment thicker than remaining segments, a little shorter than third, together
with second longer than third; second one-third of third; fourth noticeably
longer than fifth, a little shorter than third; distal antennal segments appro-

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84

ximately equal to fifth. Mandibles short, not projecting anteriorly. Width of
pronotum 1.8 times the length, anterior margin very slightly narrower than
posterior; posterior margin weakly roundly protuberant. Prosternum slightly
shorter than pronotum; length of prosternum in front of forecoxae equal to
coxal length. Intercoxal prosternal process half the coxal width, extending
beyond coxae and broadening, apex rounded. Length of propleuron twice its
width. Length of mesosternum half the midddle coxae, triangular depression
for prosternal process small. Mesepisterna almost rectangular, transverse;
mesepimera strongly broadening laterally. Length of metasternum half the
width at the posterior margin; width at posterior margin 1.7 times at anterior;
distance between middle and hind coxae noticeably longer than middle
coxae; mesepisterna not sharply broadening in anterior half. Femoral plates
of hind coxae laterally rather sharply tapering, their elongate part noticeably
longer than their width, narrower than half the coxal width. Reduced part of
the femoral plate in the form of a narrow angular tongue extending a little
beyond the metasternal corner. Abdomen narrowing from base of fourth
sternite, base of last sternite two-thirds of the abdominal base, its width 3.5

>

4
aA

Fig. 32. Karadromeus rostratus, sp. nov.; holotype PIN No. 2904/873: a—dorsal
view; b—ventral view. Karatau, Upper Jurassic.

85

times the length. Forefemora two-thirds the middle femora; middle femora
noticeably shorter than-hind femora; forefemora dilated at middle; middle
and hind femora uniformly weakly thickened along the entire length.
Foretibia almost two-thirds the femur, with deeply incised groove for clean-
ing antennae. Segments of foretarsi short and broad, only slightly longer than
their apical width. Body with small but distinct punctures.

Dimensions. Body length 7.2 mm, width 3.7 mm; elytral length 4.8 mm.

Comparison. Distinguished by the broader and shorter pronotum and
the abdomen less narrowed posteriorly.

Karadromeus latus Ponomarenko, sp. nov.
(Plate V, Photo 6; Figure 33)

Species name coined from ‘latus’ (Latin)—broad.

Holotype. No. 2384/567, PIN, reverse impression of beetle without
distal parts of antennae and legs. South Kazakhstan, Chimkent oblast,
Algabass region, south-western flank of Kashkarata river valley, Aulie area
near Mikhailovka village (Karatau-Mikhailovka site). Upper Jurassic, Kara-
bastau series.

Material. Besides holotype, an inpression of a beetle lying on its side,
specimen No. 2554/724 from the same site. Due to lateral disposition of this

ll
Ja

Fig. 33. Karadromeus latus, sp. nov.; a—holotype PIN No. 2384/567; b—
paratype PIN No. 2554/724. Karatau, Upper Jurassic.

64

65

86

specimen, the shape of pronotum cannot be determined and hence its formal
inclusion in this species cannot be considered conclusive.

Description. Head equal in length and width, narrowing anteriorly from
base; temples half the length of eyes; genae short. Antennal segments
broadening slightly from base to apex; first segment noticeably thicker than
remaining segments; first segment together with the second equal in length
to third. Labrum very weakly emarginate at apex. Mandibles rather short,
projecting alittle from under labrum. Width of pronotum 1.6 times its length;
width at its anterior margin nine-tenths that at the posterior; anterior angles
blunt; middle of posterior margin angularly projecting backward; pronotum
much narrower than elytra at shoulders. Width of metasternum at posterior
margin 2.2 times its length, anterior margin narrower than posterior; distance
between middle and hind coxae noticeably greater than the length of middie
coxae. Metepisterna strongly but not sharply broadening in anterior half.
Elongate part of femoral piates one-and-a-half times longer than its width.
Abdomen narrowing from base of third sternite, basal width of last sternite
half of abdominal base, its length one-fourth its width. All femora are weakly
clavate, their widest part at apical third approximately twice that at apex;
middle and forefemora* almost equal, slightly shorter than forefemora.
Tibiae slender, slightly broadening from base to apex; fore- and middle tibiae
slightly shorter than femora, hind tibiae equal to the corresponding femora;
apical spurs of tibiae short. Tarsi slightly shorter than tibiae, segments of
fore- and middle tarsi short, somewhat broad, segments of hind tarsi almost
linear. Body with small dense punctation.

Dimensions. Body length 8.5—9.3 mm, width 5.3 mm; elytral length
5.8-6.2 mm.

Comparison. Distinguished by narrow pronotum which is much nar-
rower than the elytra at shoulders, and the more strongly posteriorly narrow-
ing abdomen.

Karadromeus elongatus Ponomarenko, sp. nov.
(Plate V, Photo 7; Figure 34a)

Species name coined from “elongatus’ (Latin)—clongate.

Holotype. No. 3015/363, PIN, impression of a beetle without apices of
antennae and of almostall legs. Trans-Baikal, Chita oblast, Balei region, right
bank of Unda river, 2 km upstream Zhidka settlement (Unda site). Lower
Cretaceous, Aptian-Albian?, Balei series

Material. Holotype.

Description. Body of beetle elongate, cylindrical. Head noticeably
longer than its basal width; temples and genae slightly shorter than eyes.

* An obvious mispmint in the Russian original. Should read ‘hind’ instead—General Editor.

87

Fig. 34. Species of genus Karadromeus. a—K. eiongatus, sp. nov.; holotype PIN
No. 3015/363, Unda, Jurassic; b—K. mongolicus, sp. nov.; holotype PIN No. 3145/
753. Mongolia, Anda-Khuduk, Lower Cretaceous.

Second antennal segment half the third. Pronotum slightly narrowing in the
anterior fourth, its basal width 1.7 times its length; anterior margin weakly
roundly emarginate. Mesosternum short, depression for prosternal process
large. Mesepimera almost taenioid, very slightly broadening laterally.
Metasternal length one-third the width at posterior margin; width at anterior
margin half that at the posterior; distance between middle and hind coxae
one-and-a-half times longer than middle coxae. Metepisterna weakly and
gradually broadening in anterior two-thirds. Hind coxae short, weakly and
gradually tapering laterally. Elongate part of femoral plates medial, plates
mesally and laterally tapered, with an emargination at the mesai margin,
lateral tapered part almost extending up to lateral ends of coxae. Abdomen
more than one-and-a-half times longer than meso- and metathorax together,
narrowing from base of fourth sternite, sutures between basal sternites not
noticeable. Length of last sternite one-third its basal width; its base two-thirds
the width of abdominal base. Lobes of eighth true sternite small, imperfectly
teardrop-shaped; parameres asymmetrical, short, not perfectly triangular.
Hind femora slightly thickened. Body very sparsely punctate.

Dimensions. Body length 4.3 mm, width 1.9 mm; elytral length 2.9 mm.

Comparison. Distinguished from the remaining species by elongate,
narrow body, long abdomen, long last abdominal sternite, and shape of
femoral plates with a notch at mesal margin.

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88

Karadromeus mongolicus Ponomarenko, sp. nov.
(Plate V, Photo 8; Figure 34b)

Species name coined from Mongolia.

Holotype. No. 3145/753, PIN, impression of a poorly preserved beetle
with spread out elytra. Mongolia, Uver-Khangai aimak, Ushugiin-Nuru
range, western sources of Shand-god near Anda-Khuduk pit (Anda-Khuduk
site). Lower Cretaceous, Neocomian, Anda-Khuduk series.

Material. Besides holotype, impression of a poorly preserved beetle,
No. 3145/754 from the same site. It is possible that some of the isolated elytra
collected there also belong to this species.

Description. Length of head capsule noticeably less than its width; eyes
slightly longer than temples; genae approximately half the length of temples.
Width of pronotum 1.3 times its length, broadest immediately in front of
middle from where it roundly tapers anteriorly and posteriorly; anterior —
margin a little narrower than posterior. Prosternum much shorter than
pronotum, in front of forecoxae equal in length to coxae. Prosternal process
half as wide as coxae, scarcely extending beyond posterior margin of coxae,
apically rounded. Propleura narrow, more than twice as long as wide.
Mesosternum half the length of middle coxae; triangular depression for
prosternal process large, wide. Mesepisterna small, rectangular; mesepimera
long, laterally noticeably broadening. Metasternum approximately two-and-
a-half times longer than wide, very slightly narrowing right up to the anterior
margin; its anterior margin two-thirds the posterior; distance between coxae
greater than one-and-a-half times the length of the transverse middle coxae.
Metepisterna very slightly broadening almost up to the anterior margin,
[greatly] broadening only along the anterior margin itself; width at anterior
margin one-third the metepisternal length. Width of hind coxae sharply
reduced beyond mesal third; their maximum length two-and-a-half times the
medial length. Femoral plates large, width of their elongate part more than
half that of coxae, rather sharply reduced laterally. Abdomen narrowing from
beginning of fourth sternite. Hind legs short, femoral apices scarcely extend-
ing beyond lateral margins of body, femora slightly thickened, broadest part
a little distal to middle. Tibiae noticeably shorter than femora, rather thick,
almost not broadening apically. Elytra smooth; traces of two grooves notice-
able only in the sutural part. Body with small punctures.

Dimensions. Body length 6.5—7.0 mm, width 2.8-3.0 mm; elytral length
4.7-5.0 mm.

Comparison. Distinguished from other species by the long pronotum
and long, very slightly anteriorly narrowing metastemum.

Remarks. The fossils described here are very poorly preserved. Al-
though considerable chitin has been retained in the impressions, it is difficult
to study their structure precisely. Hence the interpretation of the observed

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89

structures cannot be defended with total certainty. The structure of hind coxae
is particularly unclear. At the same time, the structure of the anteriorly very
slightly broadening metepisterna makes this form very interesting, exhibiting
the characteristic tendency towards closure of middle coxal cavities found in
the present-day ground beetles, which is rare in the Mesozoic beetles.

Genus Eodromeus Ponomarenko, gen. nov.

Genus name coined from ‘eos’ (Greek)—early, and ‘dromeus’ (Greek)—
runner.

Type species. E. fasciatus sp. nov.Lower Cretaceous of Trans-Baikal.

Description. Small, flat beetles. Head including mandibles shorter than
its length*, triangular, temples and genae shorter than eyes. Antennae short,
scarcely extending beyond pronotal base, segments thick, flagellar segments
only a little longer than broad. Pronotum almost rectangular, anteriorly very
weakly emarginate, anterior angles not sharp, pronotal base slightly narrower
than elytral base. Mesosternum with distinct triangular depression for
prosternal process. Femoral plates of hind coxae emarginate at posterior and
lateral margins. Legs short, hind tibiae not longer than femora.

Species composition. One species in the Upper Jurassic of South
Kazakhstan, three species in the Lower Cretaceous of Trans-Baikal.

Comparison. Distinguished from other genera by the shape of femoral
plates of hind coxae which are emarginate at posterior margin and strongly
so laterally. In addition, differs from most representatives of the family by
the almost rectangular pronotum.

Eodromeus antiquus Ponomarenko, sp. nov.
(Plate VI, Photo 1; Figure 35)

Species name coined from ‘antiquus’ (Latin)—ancient.

Holotype. No. 2239/897, PIN, impression of an almost entire beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site); Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head with mandibles**. Maximum width of head capsule
1.7 times its length, head narrowing anteriorly from base. Eyes slightly longer
than temples, genae short. First antennal segment slightly thicker than the

*Correct as per the Russian original. Should read “width”—General Editor.

**There seems to be an omission in the Russian original. The completed phrase would have
drawn a comparison between the length of the head, including the mandibles, and its width. If
Figure 35 is drawn to scale, these two dimensions are approximately equivalent—Scientific
Editor.

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90

rest, scarcely shorter than third, one-and-a-half times longer than second;
second noticeably longer than broad. Flagellar segments commencing from
fourth almost equal, only slightly longer than second. Labrum rather deeply
emarginate at apex. Mandibles one-and-a-half times longer than wide, api-
cally incurved. Palpal segments short, rather thick. Lateral lobes of mentum
large, not shorter than their width. Gular plate narrow, its length almost three
times the medial width. Width of pronotum 1.8 times the length, anterior
margin almost the same length as posterior, with very slight trapezoidal
emargination, posterior margin weakly, roundly protuberant. Pronotum
broadest in the anterior third. Prosternum slightly shorter than pronotum,
anterior to forecoxae its length equal to that of forecoxae. Width of prosternal
process half that of coxae, almost not extending beyond them. Length of
propleuron twice the width. Mesosternum rather long, not shorter than
middle coxae, triangular depression narrow. Mesepisterna almost rectan-
gular, mesepimera very slightly broadening laterally. Middle coxae converg-
ing, a little longer than broad. Metasternal width at posterior margin 3.3 times
its length, width at posterior margin 2.2 times that at the anterior. Distance
between middle and hind coxae equal to length of middle coxae. Transverse
metasternal suture almost straight. Hind coxae from middle almost not
tapered laterally, lateral margin half the medial. Length of femoral plates
sharply reduced from middle laterally; rather wide, angular lateral process
along the anterior coxal margin almost reaching the corners of metasternum.
Emargination on posterior margin of femoral plates wide and shallow, lateral
emargination deep. Abdomen slightly broadening from base to the third
sternite, and tapering beyond; width of last sternite two-thirds the abdominal
width*, last sternite one-third as long as wide. Femora gradually dilating,
clavate, widest in apical third. Forefemora noticeably shorter than middle
femora, and two-thirds the hind femora. Tibiae rather thick, middle and hind
tibiae noticeably broadening in apical fourth, hind tibiae weakly broadening
from base, not broadening before apex. Fore- and middle tibiae equal in
length, hind tibiae 1.7 times longer. Groove on the foretibiae extending up
to their middle. Spurs on fore- and middle tibiae much shorter than the first
tarsal segment, those on the hind tibiae not shorter than it [first tarsal
segment]. Tarsal segments rather broad, slightly broadening from base to
apex, hind tarsi slightly shorter than tibiae, their segments almost equal in
length.

Dimensions. Body length 7.2 mm, width 3.0 mm; elytral length 4.7 mm.

Comparison. Distinguished by short, narrow prosternal process and
shape of the femoral plates of hind coxae. which are highly extended
laterally.

*Possibly referring to the basal width (cf. page 92)—General Editor.

91

Fig. 35. Eodromeus antiquus, sp. nov.; holotype PIN No. 2239/897: a—dorsal view;
b—ventral view. Karatau, Upper Jurassic.

Eodromeus sternalis Ponomarenko, sp. nov.
(Plate VI, Photo 2; Figure 36)

Species name coined from ‘sternon’ (Greek)—breast.

Holotype. No. 3191/5, PIN, impression of beetle without legs and an-
tennae. Trans-Baikal, Chita oblast, Krasnochikoi region, well No. 102 (Chita
Geological Administration, 1963), depth 138.2 m. Neocomian, Tignin series.

Material. Holotype.

Description. Head short, its length together with mandibles two-thirds
the width at posterior margin, narrowing anteriorly from base. Length of
temples and genae half that of eyes. Length of gular plate one-and-a-half
times the medial width. Width of pronotum almost twice the length, its

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92

maximum width a little anterior middle. Prosternum scarcely shorter than
pronotum, anterior to forecoxae one-and-a-half times the length [of
forecoxae]. Prosternal process large, only slightly narrower than forecoxae,
projecting far beyond their posterior margin. Mesosternum slightly shorter
than middle coxae, triangular depression wide. Mesepimera long, almost not
laterally broadening. Middle coxae a little longer than wide, metasternal
process between them only slightly narrower than coxae. Length of metaster-
num one-third the width, width at posterior margin 2.3 times that at anterior;
distance between middle and hind coxae very slightly more than the length
of middle coxae. Length of hind coxae sharply reduced laterally from medial
fourth, further tapering gradually toward lateral margin. Length of femoral
plates sharply reduced before middle of coxae [=just inside of mesal half],
almost not extending laterally along the anterior margin; lateral and posterior
margins very weakly emarginate. Abdomen narrowing from base of fourth
sternite, width of last sternite half the abdominal base, its width 3.5 times the
length. Elytra with weak but distinct grooves. Integument almost smooth,
with sparse fine punctation.

Dimensions. Body length 7.1 mm, width 3.6 mm; elytral length 4.6 mm.

Comparison. Differs from other species in long prosternum, and
posteriorly and laterally weakly emarginate femoral plates. _

Se ee

Wn
|

a (eee) b

Fig. 36. Eodromeus sternalis, sp. nov.; holotype PIN No. 3191/5: a—dorsal view;
b—ventral view . Chikoi Basin, Lower Cretaceous.

93

Eodromeus dissectus Ponomarenko, sp. nov.
(Plate VI, Photos 3, 4; Figure 37)

Species name coined from ‘dissectus’ (Latin)—cut.

Holotype. No. 1989/2967, PIN, impression of a beetle without head and
pronotum. Trans-Baikal, Buryat ASSR, Eravnin region, left bank of Vitim
river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian- Goteriv, Zazin series.

Material. Besides holotype, one impression of beetle without one anten-
na and major part of legs, specimen No. 3064/863* from the same site. Only
the dorsal structure distinctly visible in the impression.

Description. Length of head with mandibles slightly less than its width,
width of head capsule 1.7 times its length, narrowing from base anteriorly.
Eyes very slightly shorter than temples, genae less than half the length of
eyes. Antennae projecting beyond pronotal base by four segments; first

Fig. 37. Eodromeus dissectus, sp. nov.; a—holotype PIN No. 1989/2967;
b—paratype PIN No. 3064/803. Baisa, Lower Cretaceous.

*3064/803 in Figure legend—General Editor.

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94

segment slightly thicker than flagellar segments, almost twice the length of
second; third one-third longer than second; fourth and following segments
subequal, two-thirds the third, only the last slightly longer. Labrum very
slightly emarginate. Width of pronotum almost twice the length, anteriorly
with shallow straight emargination; pronotum rounded laterally, narrowing
anteriorly and posteriorly from middle, slightly protuberant at posterior
margin. Anterior angles of pronotum drawn out posteriorly, blunt anteriorly.
Disk of pronotum with weak transverse oval raised portion, divided by flat
longitudinal groove, more distinct in posterior half. Flat lateral flange of
pronotum narrow, [extreme] lateral margin not margined. Prosternum
noticeably shorter than pronotum; anterior to forecoxae not shorter than
them. Mesosternum rather long, only slightly shorter than middle coxae,
triangular depression narrow. Length of mesopleura not less than width,
pleural suture slightly oblique so that mesepisterna and mesepimera almost
rectangular; mesepimera long, more than half of mesepisterna. Middle coxae
round, converging, metasternal process between them less than half the coxal
width. Width of metasternum at posterior margin almost three times the
length posterior margin; posterior margin 2.3 times the anterior. Distance
between middle and hind coxae one-and-a-half times longer than middle
coxae. Longitudinal lines apparently corresponding to internal keels
proceeding anteriorly along the metasternum from middle coxae backward
and from hind coxae approximately to their lateral third. Metepisternum in
anterior half strongly but not sharply broadened, width at anterior margin
two-fifths the length. Length of hind coxae sharply reduced laterally from
medial third, more laterally maintaining the same length right up to the lateral
margin, length before lateral margin only two-thirds the maximum length.
Femoral plates of hind coxae small, noticeably narrower than half of coxa,
on anterior margin with small angular lateral projection, by far not extending
to the corners of metasternum; lateral margin in anterior half with a deep
emargination, roundly projecting posteriorly from it, posterior margin with
a narrow and rather deep notch, postero-lateral corners somewhat extended,
rounded. Abdomen weakly narrowing posteriorly from the base of third
sternite; abdominal base 1.7 times as wide as last sternite; last sternite
triangular, its width 1.4 times the length. Fore- and middle femora broadened
uniformly, reaching maximum width almost at middle, hind femora gently
clavate,widest in apical third, middle femora only slightly shorter than hind
femora. Tibiae rather thick, noticeably broadening from base to apex, their
outer margin with spines, length of foretibiae nearly half and middle tibiae
noticeably shorter than hind tibiae; segments of foretarsi broadening, trian-
gular, length not greater than apical width. Spurs of middle and hind tibia
shorter than first tarsal segment. Middle tarsi equal to tibiae in length, basal
segments broadening from base to apex, distal ones slender, almost virgate;
first and last segments equal, almost twice as long as the remaining. Hind

71

95

tarsi shorter than tibiae, their segments short, first segment one-and-a-half
times longer than second. Elytra smooth. Body without distinct punctation
dorsally, with sparse and small punctures ventrally.

Dimensions. Body length 6.0—7.5 mm, width 2.7—3.1 mm; elytral length
4.0-5.0 mm.

Comparison. Distinguished by longer metasternum and small short
femoral plates with deep emarginations and extended postero-lateral corners.

Eodromeus major Ponomarenko, sp. nov.
(Plate VI, Photo 5; Figure 38)

Species name coined from ‘major’ (Latin)— large.

Holotype. No. 3064/864, PIN, impression of beetle without antennae
and large part of legs. Trans-Baikal, Buryat ASSR, Eravnin region, left bank
of Vitim river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Width of head at posterior margin 1.6 times the length of
head with mandibles and 2.5 times the length of head capsule, narrowing
from base anteriorly. Temples two-thirds the length of eyes, genae very short.
Mandibles approximately twice as long as wide, apices incurved. Lateral

Fig. 38. Eodromeus major, sp. nov.; holotype PIN No. 3064/864. Basia, Lower
Cretaceous.

96

lobes of mentum longer than wide, apically rounded. Gular plate broadening
in apical third, twice as long as maximum width anterior to middle; anterior
margin slightly narrower than posterior, anterior corners slightly drawn out.
Prosternum slightly shorter than pronotum, anterior to forecoxae scarcely
shorter than slightly transverse coxae. Prosternal process half as wide as
coxae, extending slightly beyond them, rounded apically. Propleura 2.2 times
longer than wide. Mesosternum equal in length to middle coxae, triangular
depression equilateral. Mesopleuron distinctly less long than wide, pleural
suture intersecting almost diagonally, mesepimeron strongly broadening
laterally. Middle coxae oval, transverse, convergent; intercoxal metasternal
process very small. Length of metasternum one-third its width at posterior
margin, width at anterior margin half of posterior. Distance between middle
and hind coxae one-third longer than middle coxae. Metepisternum in
anterior half more strongly but not sharply broadening, its length two-and-
a-half times the width. Elongate part of hind coxae narrow, not more than
one-fourth their width, more laterally the coxae gradually tapering toward
the lateral margin. Femoral plates of hind coxae small, at anterior margin
with narrow lateral angular projection, by far not extending to metasternal
corners lateral margin before the middle with deep angular emargination,
posterior margin with deep emargination somewhat shifted medially from
middle of posterior margin, postero-lateral corner broadly rounded. Ab-
domen rather strongly narrowing from the base of second sternite; width of
last sternite half of abdominal base, triangular, its width 2.8 times its length.
Legs short, middle femora scarcely extending beyond body margins, slightly
thickened, their maximum width somewhat distal to middle. Tibiae equal in
length to femora, strongly broadening in distal half, outer margin with spines,
spurs not shorter than first tarsal segment. The latter almost twice as long as
second segment. Metasternum with dense, small punctures, punctation of
other sclerites indistinct.

Dimensions. Body length 11.3 mm, width 6.0 mm; elytral length 7.5
mm.

Comparison. Distinguished from other species by short head. Resem-
bles E. sternalis in general body shape and short head, but differs in the
smaller prosternal process and shape of hind coxae; resembles E. dissectus
in the structure of hind coxae, but differs in the shorter head and metasternum.

Family CARABIDAE Latreille, 1802

Subfamily Protorabinae Ponomarenko, Subfam, Nov.

Diagnosis. Clypeus not extending up to antennal socket. Forecoxal cavities
open. Lateral walls of middle coxal cavities formed by mesosternum,

97

72 mesepisterna, mesepimera and metasternum. Hind coxae not separating
metapleuron and abdomen. Both spurs of foretibiae apical.
Composition. Five genera in the Jurassic of Soviet Central Asia and
South Kazakhstan, and the Lower Cretaceous of Trans-Baikal.
Comparison. Distinguished from all other carabids by the inclusion of
metepisterna into lateral walls of middle coxal cavities.

Genus Protorabus Ponomarenko, gen. nov.

Genus name coined from ‘protos’ (Greek)—first, and from genus Carabus.
Type species. P. planus sp. nov. Upper Jurassic of South Kazakhstan.
Description. Medium-sized, broad, flat beetles. Head triangular, trans-

verse. Genae shorter than temples. Antennae rather long, their segments

slightly broadening apically. Prosternum transverse, its maximum width
anterior to middle, constricted* behind middle. Mesosternum and distance
between middle and hind coxae not longer than middle coxae. Hind coxae
oblique. Femoral plates laterally reaching only up to coxal middle. Abdomen
short, sternites movable, apex blunt. Base of last sternite only slightly
narrower than abdominal base. Legs short, femora scarcely extending beyond
lateral margins of body. Elytra with grooves.

Species composition. Three species in the Upper Jurassic of South

Kazakhstan.

Comparison. Distinguished by short, posteriorly weakly tapering
abdomen with broad last sternite.

Protorabus planus Ponomarenko, sp. nov.
(Plate VI, Photos 6, 7; Figure 39)

Species name coined from ‘planus’ (Latin)—flat.

Holotype. No. 2066/3185, PIN, impression of a beetle without antennae
and large part of legs. South Kazakhstan, Chimkent oblast, Algabass region,
south-western flank of Kashkarata river valley, Aulie area near Mikhailovka
village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Besides holotype, nearly entire beetles, PIN Nos. 2904/928,
2066/3187 and metathorax with abdomen, No. 2784/1299 from the same site.

Description. Head transverse, triangular, head capsule almost half its
width at posterior margin, uniformly narrowing anieriorly from base.
Temples shorter than eyes and longer than genae. Labrum weakly emarginate
at apex. Mandibles almost equal to head capsule in length. Submental lobes

*The Russian original reads ‘behind middle prostemum’, which makes no sense. A com-
parison with the generic descriptions of Cordorabus and Cretorabus on subsequent pages,
suggests that the word ’perednegrad’ (= prostemum) was accidentally substituted for the word
“‘peretyanuta’ ( = constricted)—Scientific Editor.

98
narrow, incision between them rather deep. Antennae projecting far beyond

pronotal base, their first segment noticeably thickened, second equal in
length and width, third equal to first and second together, remaining flagellar

Reon

on \
AAW,
1 |

Cc

73 Fig. 39. Protorabus planus, sp. nov.; a, b—holotype PIN No. 2066/3185:
a—dorsal view; b—veniral view; c, d—paratype PIN No. 2904/928: c—dorsal
view, d—ventral view. Karatau, Upper Jurassic.

73

74

29

segments-equal, two-thirds the length of third. Pronotum noticeably longer
than head, its length two-thirds the basal width, anterior margin with shallow
rounded emargination. Pronotum margined laterally, posterior margin with
longitudinal groove. Prosternum noticeably shorter than pronotum, anterior
to forecoxae approximately as long as forecoxae. Prosternal process half as
wide as coxae straight, short, scarcely extending beyond coxae. Propleura
almost twice as long as wide. Mesosternum shorter than middle coxae, with
triangular depression for prosternal process, mesepisterna almost square,
mesepimera short, laterally almost not widening. Lengih of metasternum
approximately one-fourth the width at posterior margin, anterior margin
approximately haif the width of posterior; distance between [middle and
hind] coxae noticeably shorter than middle coxae. Posterior margin of
metasternum angularly projecting, distance from apex of angle to transverse
metasternal suture only slightly shorter than hind coxae. Hind coxae laterally
tapering, triangular, femoral plates wider than long, angularly extending
posteriorly and laterally. Metepisterna sharply broadening in front of anterior
margin, width at anterior margin half the length, posterior margin rather wide;
metepimera mesally not broadening. In impressions, abdomen only slightly
longer than meso- and metathorax together, narrowing from base of penul-
timate sternite, length of last sternite one-third the basal width. Femora not
strongly but uniformly thickened, tibiae flat, broadening from base to apex.
Foretibiae slightly shorter than femora, on inner side with groove for cleaning
antennae and small notch preapically. Apical spurs shorter than primary
tarsal segments. Four basal segments of foretarsi broadening toward apex,
scarcely longer than their width. Hind tibiae almost linear, broadening only
preapically, not shorter than femora. Furrows of elytra wide, flat, indistinct.
Integument with small, sparse punctation.

Dimensions. Body length 14.7-16.0 mm, width 6.7—8.3 mm; elytral
length 8.6-9.7 mm.

Comparison. Distinguished by longer pronotum and more sharply
anteriorly narrowing metastenum.

Protorabus nigrimonticola Ponomarenko sp. nov.
(Plate VI, Photo 8; Figure 40)

Species name coined from ‘nigrimonticola’ (Latin)—inhabitant of Black
mountains (Karatau).

Holotype. No. 2997/1851, PIN, impression of an almost entire beetle.
South Kazakhstan, Chimkent obiast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

100

Material. Besides holotype, one impression of beetie without legs, No.
2997/485, two impressions of metathorax and abdomen, specimen Nos.
2997/1866 and 2066/2531 from the same site.

Description. Head including mandibles almost equal in length and
width, triangular, head capsule two-thirds its basal width, uniformly narrow-
ing from base anteriorly. Temples less than two-thirds the eyes, genae shorter
than temples. Apex of labrum truncated. Submental lobes rather broad,
anteriorly rounded. Antennae projecting far beyond pronotal base; their first
segment thickened, equal in length to second and third together; second
small, almost equal in length and width, half the length of third; third
noticeably longer than fourth; remaining flagellar segments equal in length.
Pronotum scarcely longer than head, almost half the basal width, anterior
margin with rather deep rounded emargination, laterally widely margined.
Prosternum much shorter than pronotum, prosternum anterior to forecoxae
shorter than them. Prosternal process half the forecoxae in width, straight,
noticeably extending beyond their posterior margin. Mesosternum shorter
than middle coxae, anteriorly with triangular depression for prosternal
process, mesepisterna transverse, mesepimera short, noticeably broadening
laterally. Metasternum almost one-third the width of posterior margin,
anterior margin half the width of posterior, distance between middle and hind

Fig 40. Protorabus nigrimonticola, sp. nov.; holotype PIN No. 2997/1851: a—
dorsal view; b—ventral view. Karatau, Upper Jurassic.

75

101

coxae slightly longer than midcoxae. Hind margin of metasternum and
transverse metasternal suture angularly extending posteriorly, distance be-
tween them much shorter than hind coxae. Hind coxae triangular, femoral
plates almost equal in length and width, coxae 2.3 times longer than femoral
plates. Metepisterna gradually broadening anteriorly almost from posterior
margin, width at anterior margin half the length. Abdomen noticeably longer
than meso- and metathorax together, narrowing posteriorly almost from base,
length of last sternite three-fourths its basal width. Femora weakly but
uniformly thick, foretibiae shorter than femora, almost not broadening
toward apex, inner side with groove and apical third with notch for cleaning
antennae. Hind tibiae slightly broadening from base to apex; hind tarsi shorter
than tibiae, their first segment longer, almost equal to third and fourth
together; second noticeably shorter than first*, equal to fifth segment. Integu-
ment with fine, sparse punctation. Grooves on elytra rather deep and sharp.

Dimensions. Body length 7.6—8.8 mm, width 3.8-4.6 mm; elytral length
5.2-6.0 mm.

Comparison. Distinguished by shorter pronotum, longer and less sharp-
ly anteriorly narrowing metasternum, and angular transverse metasternal
suture.

Protorabus magnus, Ponomarenko, sp. nov.
(Plate VI, Photo 9; Figure 41)

Species name coined from ‘magnus’ (Latin)—large.

Holotype. No. 2554/447, PIN, impression of metasternum, abdomen
and hind legs of beetle. South Kazakhstan, Chimkent oblast, Algabass region,
south-western flank of Kashkarata river valley, Aulie area near Mikhailovka
village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Length of mestasternum two-fifths the width at posterior
margin, width at posterior margin 1.7 times that at anterior. Posterior margin
of metasternum angularly projecting backward, transverse metasternal suture
straight; distance between apex of angle and transverse suture half the length
of hind coxae. Hind coxae triangular, femoral plates equal in length and
width, half the coxal width, laterally strongly emarginate. Metepisterna
sharply broadening in anterior third, angularly tapering at posterior margin.
Abdomen narrowing from fourth sternite, length of last sternite one-third the
basal width. Hind femora clavate, widest in distal third, tibiae linear, equal
to femora in length, tarsi noticeably shorter, first segment one-and-a-half
times longer than second, second to fourth equal, noticeably broadening

*In the Russian original words from tibiae (line 2 from top) to ‘shorter than’ (line 4 top)
repeated, which have been deleted here—General Editor.

76

102

Fig. 41.* Protorabus magnus sp. nov.; holotype PIN No. 2554/447. Karatau,
Upper Jurassic.

apically, last equal to the preceding two together. Integument with small,
dense punctation.

Dimensions. Length of fragment 15 mm, width 11 mm; length of beetle
apparently about 20 mm.

Comparison. Distinguished by long metasternum, posteriorly narrow-
ing metepisterna and long terminal abdominal sternite.

Genus Ovrabites Ponomarenko, gen. nov.

Genus name coined from ‘ovum’ (Latin)—egg.

Type species. O. ovalis sp. nov. Upper Jurassic of South Kazakhstan.

Description. Small, oval, flat beetles. Head triangular, approximately
equal in length and width. Antennae long, their segments slightly apically
broadening. Prosternum transverse, trapezoidal, maximum width at base.
Prosternal process broadening toward apex, apically truncate. Mesosternum
with a triangular depression for prosternal process. Hind coxae oblique,
length of femoral plates reduced laterally, plates much narrower than hind
coxae. Abdomen rather long, posteriorly narrowing almost from base; base
of the last abdominal sternite much narrower than remaining sternites, width
half the abdominal base, apex of abdomen rather pointed. Legs short, femora
scarcely extending beyond body margins. Elytra smooth.

Species composition. Two species in the Late Jurassic of South Kaza-
khstan.

*This figure was printed upside down in the Russian original—General Editor.

103

Comparison. Differs from all other genera in the shape of pronotum—
widest at base and uniformly tapering anteriorly.

Ovrabites ovalis Ponomarenko, sp. nov.
(Plate VII, Photo 1; Figure 42)

Species name coined from ‘ovalis’ (Latin)—egg-shaped.

Holotype. No. 2904/872, PIN, impression of beetle without large part of
legs; South Kazakhstan, Chimkent oblast, Algabass region, south-western
flank of Kashkarata river valley, Aulie area near Mikhailovka village
(Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

eo
eee

ai

j

/

Fig. 42. Ovrabites ovalis, sp. nov.; holotype PIN No. 2904/872: a—ventral view;
b—dorsal view of head, and pronotum. Karatau, Upper Jurassic.

UY

104

Material. Holotype and almost entire impressions of beetles, specimen
Nos. 2384/554, 2904/875, 2554/526, 2784/1326 from the same site.

Description. Head including mandibles very slightly shorter than its
basal width, uniformly tapering from base anteriorly. Temple and genae very
short, much shorter than eyes. Labrum truncate. Mandibles half the length
of head capsule. Submental lobes rather wide, apically rounded. Antennae
projecting by one-third beyond pronotal base. Their first segment thicker
than remaining segments, half of third; second cylindrical, equal in length
and width, half the length of third; third and fourth slightly longer than the
remaining flagellar segments. Pronotum almost same length as head, ap-
proximately half its basal width; anterior margin with deep, almost rectan-
gular emargination; posterior margin weakly angularly projecting backward;
laterally margined. Prosternum only slightly shorter than pronotum, anterior
to forecoxae noticeably longer than them. Prosternal process slightly nar-
rower than coxae. Propleura twice as long as wide. Mesosternum very short,
almost taenioid, half the length of middle coxae. Length of metasternum
one-third its width at posterior margin; width at posterior margin 1.8 times
that at the anterior; distance between middle and hind coxae almost equal to
length of middle coxae. Posterior margin of metasternum angularly project-
ing, transverse metasternal suture posteriorly weakly convex, distance be-
tween it and apex of angle at hind margin less than half the length of hind
coxae. Hind coxae nearly same length, noticeably shorter than femoral plates.
Femoral plates longer than width at anterior margin, basal margin emar-
ginate, postero-lateral angles acute, extended. Metepisterna sharply broaden-
ing in front of anterior end. Abdomen twice as long as meso- and metathorax
together, narrowing almost from base; last sternite very slightly longer than
penultimate, its basal width 3.5 times the length. Fore- and middle femora
slightly thickened, hind femora considerably thickened. Tibiae slender,
scarcely broadening toward apex, middle tibiae shorter than femora, hind
tibiae very slightly longer. Hind tarsi slender, linear, their segments almost
equal in length, scarcely broadening toward apex. Integument with fine and
rather dense punctation.

Dimensions. Body length 6.2—7.2 mm, width 2.7—3.0 mm; elytral length
4,.0-4.8 mm.

Comparison. Distinguished by shorter metastemum.

Ovrabites jurassicus Ponomarenko, sp. nov.
(Plate VII, Photo 2; Figure 43)

Holotype. No. 2904/877, PIN, impression of beetle without legs and apices
of antennae. South Kazakhstan, Chimkent oblast, Algabass region, south-
western flank of Kashkarata river valley, Aulie area near Mikhailovka village
(Karatau-Mikhailovka site). Upper Jurassic, Karabastau series.

105

Material. Besides holotype, impression of beetles without legs, speci-
men No. 2997/407 from the same site, and No. 2452/33 from the Karatau-
Galkino site.

Description. Head triangular, noticeably shorter than its basal width,
narrowing from posterior margin of eyes anteriorly. Temples noticeably
longer than genae, two-thirds of eyes. Labrum truncate. Mandibles short, half
the length of head capsule. Submental lobes small, apically rounded. Anten-
nae extending beyond pronotal base by half their length. Their third segment
is equal to the first and second together, scarcely thinner than the first; fourth
noticeably shorter than third; remaining flagellar segments equal to fourth
segment. Pronotum noticeably longer than head, half the basal width; anterior
margin with rather deep rounded emargination, posterior margin almost
straight, prosternum slightly shorter than pronotum, anterior to forecoxae
shorter than them. Prosternal process scarcely projecting beyond coxae,
slightly narrower than them. Length of propleura two-thirds the maximum
width. Mesosternum two-thirds the length of middle coxae. Length of
metasternum two-fifths the basal width, width at anterior margin ap-

78 proximately half that posterior. Distance between middle and hind coxae
much longer than middle coxae. Posterior margin of metasternum angular,
transverse metasternal suture roundly projecting posteriorly. Distance be-
tween them half the hind coxae. Length of hind coxae sharply reduced
laterally from medial third, length gradually decreasing further on, their

—"

a

Fig. 43. Ovrabites jurassicus, sp. nov.; a, b—holotype PIN No. 2904/877: a—dor-
sal view, b—ventral view; c—paratype PIN No. 2452/83. Karatau, Upper Jurassic.

TE)

106

raised part just slightly less than femoral plate in length. Length of femoral
plate equal to width at anterior margin, lateral margin strongly emarginate,
postero-lateral angles acute, extended. Metepisterna broadening from middle
anteriorly. Abdomen one-and-a-half times longer than meso- and metathorax
together, tapering almost from base; last sternite not longer than penultimate,
almost one-fourth the basal width. Integument with fine, rather dense puncta-
tion.

Dimensions. Body length 5.9-6.4 mm, width 3.0-3.2 mm; elytral length
4.24.4 mm.

Comparison. Distinguished by longer metasternum, less sharply nar-
rowing anteriorly.

Genus Cordorabus Ponomarenko, gen. nov.

Genus name coined from ‘cor’ (Latin)—heart.

Type species. C. notatus sp. nov. Upper Jurassic of South Kine

Description. Small, cylindrical beetles. Head triangular, transverse.
Eyes somewhat shifted dorsally. Antennae slender, projecting a little beyond
pronotal base, their segments almost not apically broadening. Prosternum
transverse, widest anterior to middle, behind it slighly constricted. Prosternal
process noticeably narrower than forecoxae. Mesosternum half the length of
middle coxae, with triangular depression for prosternal process at middle.
Hind coxae, oblique, femoral plates laterally reduced, scarcely projecting
into lateral half of coxa, laterally not emarginate. Abdomen rather long,
posteriorly tapering almost from base, apex pointed. Legs short, femora
scarcely extending beyond lateral body margins. Tarsi long, slender. Elytra
smooth.

Species composition. Three species in the Late Jurassic of South
Kazakhstan.

Comparison. Resembles Protorabus in the shape of pronotum, but
differs in pointed abdomen and smooth elytra.

Cordorabus notatus Ponomarenko, sp. nov.
(Plate VII, Photo 3; Figure 44)

Species name coined from ‘noton’ (Greek)—back.

Holotype. No. 2066/2739, PIN, impression of a beetle without abdomen.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype and almost entire impressions of beetles, PIN Nos.
2066/2879, 2784/1320, 2997/534, 2997/1867 from the same site.

80

107

Description. Head noticeably shorter than its basal width, anteriorly
narrowing, retracted under pronotum almost up to eyes. Temples slightly
longer than genae, almost equal to eyes in length. Labrum apically emar-
ginate. Mandibles rather short, blunt, two-thirds the length of head capsule.
Antennae extending beyond pronotal base by only three segments. First
antennal segment shorter and almost not thicker than third, third 1.3 times
longer than fourth, remaining flagellar segments almost equal. Pronotum
longer than head, length two-thirds the width at posterior margin, anterior
margin with shallow rounded emargination, posterior margin straight.
Prosternum approximately half the length of pronotum, in front of forecoxae
slightly shorier than them. Prosternal process almost not extending beyond
forecoxae. Length of propleuron twice its maximum width. Width of
metasternum at posterior margin 3.5 times its length, width at anterior margin
half that at posterior. Distance between middle and hind coxae scarcely

Fig. 44. Cordorabus notatus, sp. nov.; holotype PIN No. 2066/2739: a—ventral
view; b—dorsal view of head, and pronotum. Upper Jurassic.

108

longer than middle coxae. Posterior margin of metasternum angular,
transverse metasternal suture roundly projecting posteriorly; distance be-
tween the angle and transverse suture one-third the length of hind coxae.
Hind coxae laterally reduced [tapering] right up to lateral margins, ap-
80 proximately half the length of femoral plates, their posterior margins forming
rather smooth curve. Length of femoral plates equal to width at anterior
margin, anteriorly and at postero-lateral corners extended laterally, lateral
margin slightly protuberant medially. Metepisterna sharply broadening in

Fig. 45. Representatives of genus Cordorabus: a—C. antennatus, sp. nov.; holotype
PIN No. 2066/2363; b—C. minimus, sp. nov.; holotype PIN No. 2239/905. Karatau,

Upper Jurassic.

109

front of anterior margin. Fore- and middle femora uniformly thickened, hind
femora clavate; fore- and middle femora almost equal in length, hind femora
slightly longer. Foretibiae noticeably and middle tibiae slightly shorter than
but hind tibiae equal to [corresponding] femora. Foretibiae with longitudinal
groove for cleaning antennae, apical spurs stiff, slightly shorter than first
tarsal segment. First segment of hind tarsus longer than remaining segments,
fourth shortest. Integument with fine, dense punctation.

Dimensions. Body length 6.1-6.5 mm, width 2.7—3.0 mm; elytral length
3.84.2 mm.

Comparison. Distinguished by shorter antennae, shorter prosternum and
head, and shape of femoral plates.

Cordorabus antennatus Ponomarenko, sp. nov.
(Plate VII, Photo 4; Figure 45a)

Species name coined from ‘antenna’ (Latin)—antenna, feeler.

Holotype. No. 2066/2363, PIN, impression of a beetle without abdomen.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

81 Material. Holotype.

Description. Head scarcely shorter than its basal width, nasil up to
eyes under pronotum. Temples slightly shorter than eyes, genae very short,
antennae attached near anterior margin of eyes. Antennae projecting beyond
pronotal base by four segments. First antennal segment thickened, not longer
than fourth, shorter than third which is longer than first and second together;
third longer than fourth; remaining flagellar segments almost equal.
Pronotum very slightly longer than head, approximately half the basal width,
anterior margin with shallow, almost rectangular emargination. Prosternum
anterior to forecoxae slightly longer than them, prosternal process much
narrower than forecoxae, noticeably broadening posteriorly. Length of
propleuron two-and-a-half times its maximum width. Length of metasternum
one-third its basal width, posterior margin 1.7 times of anterior. Distance
between middle and hind coxae very slightly larger than diameter of the
former. Metepisterna sharply broadening in front of anterior margin. Hind
coxae rather sharply tapered beyond medial fourth, further on almost un-
tapered right up to lateral margins. Femoral plates slightly shorter than width
at anterior margin, markedly extending laterally beyond anterior margin,
postero-lateral corners almost right-angled, lateral margin scarcely distinctly
protuberant. Fore- and middle femora equal in length, hind femora one-third
longer, fore-femora uniformly thickened almost along entire length, middle
femora thickest just slightly distal to middle, and hind femora slightly
proximal to middle. Middle and hind tibiae equal in length to femora, not

82

110

strongly broadening from base to apex, apical spurs much shorter than basal
tarsal segment. Hind tibiae equal in length to four hind tarsal segments, first
two segments slightly longer than the subequal remaining segments. Body
with rather fine punctation, punctures noticeably larger only on metasternum.
Dimensions. Body length 9.4 mm, width 4.3 mm; elytral length 6.2 mm.
Comparison. Distinguished by longer antennae, prosternum and head,
and shape of femoral plates.

Cordorabus minimus Ponomarenko, sp. nov.
(Plate VII, Photo 5; Figure 45b)

Species name coined from ‘minimus’ (Latin)—smallest.

Holotype. No. 2239/905, PIN, impression of a beetle without antennae
and legs, with unfolded wing. South Kazakhstan, Chimkent oblast, Algabass
region, south-western flank of Kashkarata river valley, Aulie area near
Mikhailovka village (Karatau-Mikhailovka site). Upper Jurassic, Karabastau
series.

Material. Holotype.

Description. Head slightly shorter than its basal width. Temples slightly
shorter than eyes, genae half their length. Pronotum equal to head in length,
its basal width 1.3 times the length, widest in anterior third, posteriorly
roundly narrowing, not broadening in front of posterior corners, anterior
margin with shallow rounded emargination. Length of metasternum ap-
proximately one-third the width at posterior margin, width at anterior margin
half of that of posterior; distance between middle and hind coxae slightly
longer than the former. Metepisterna gradually broadening anteriorly from
middle. Hind coxae laterally short*. Femoral plates equal in length and width.
Wing with very large pterostigma, “oblongum” rather wide at anterior
margin. Body with fine sparse punctation.

Dimensions. Body length 4.8 mm, width 2.8 mm; elytral length 3.2 mm.

Comparison. Differs from other species in long narrow pronotum which
is narrowest at posterior margin.

Remarks. The species is described froma single, rather poorly preserved
specimen, and hence its inclusion in this genus is not wholly reliable.

Genus Lithorabus Ponomarenko, gen. nov.
Genus name coined from ‘lithos’ (Greek)—stone, and genus Carabus.

Type species. L. incertus, sp. nov. Lower Jurassic of Soviet Central
Asia.

*So given in the Russian original. By context it should read ‘laterally reduced’—General
Editor.

GUL

Diagnosis. Small, flat beetles. Metasternum longer than middle coxae.
Hind coxae almost transverse, their anterior margin somewhat slanting
backward only mesally. Length of femoral plates laterally reduced but
extending to lateral margins of coxae, their midcoxal length is not less than
half the maximum length. Elytra with shallow flat grooves.

Species composition. Monotype genus.

Comparison. Distinguished from other genera by the shape of femoral
plates which are very slightly reduced laterally. Differs from Cretorabus, in
which the femoral plates also extend to lateral margins of coxae, but blunt
posterior corner of metepisternum without mesal process.

Lithorabus incertus Ponomarenko, sp. nov.
(Plate VII, Photo 6; Figure 46)

Species name coined from ‘incertus’ (Latin)—uncertain.

Holotype. No. 371/49, PIN, impression of metasternum and elytra of
beetle, Kirgiz SSR, Tonsk region, southern shore of Issyk-Kyl’ lake, cost of
Kazhdi-Sai settlement (Issyk-Kul’ site). Lower Jurassic, Dzhil’ series.

Material. Holotype.

Description. Middle coxae large, contiguous, slightly elongate. Length
of metasternum two-fifths the width of posterior margin, width at posterior
margin 1.7 times that of anterior. Distance between middle and hind coxae
slightly longer than the former. Posterior margin of metasternum extending
backward at middle as a sharp tongue-like process; transverse metasternal

&) C

Fig. 46. Lithorabus incertus, sp. nov.; holotype PIN No. 371/49. Issyk-Kul’.
Lower Jurassic.

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112

suture angularly projecting backward. Width of hind coxae 1.8 times the
length, sharply reduced beyond mesal third, and gradually reduced further
laterally. Femoral plates slightly shorter than wide, reduced near coxal
middle by approximately half, and gradually reduced more laterally, longer
than coxa at lateral margin, postero-lateral corners of the elongate part of
femoral plates noticeably extending. Metepisterna broadening anteriorly
from posterior margin, their posterior margin truncate. Grooves on elytra
indistinct, those near the suture proceeding along the suture up to elytral apex,
more lateral grooves reduced in length. Metasternum almost smooth.

Dimensions. Body length about 5 mm, width 1.8 mm; elytral length 3.1
mm.

Genus Cretorabus Ponomarenko, gen. nov.

Genus name coined from ‘creta’ (Latin)—chalk, and genus Carabus.

Type species. C. capitatus, sp. nov., Lower Cretaceous of Trans-Baikal.

Description. Small and medium-sized, broad, flattened beetles. Head
large, strongly transverse. Eyes convex, genae very short, antennae rather
short, with thick segments. Pronotum transverse, maximum width anterior
to middle, constricted behind middle. Meso- and metasterna long, latter much
longer than middle coxae. Metepisterna posteriorly tapering. Hind coxae
almost straight, their anterior margin slanted backward only in middle part
of the body. Length of femoral plates sharply reduced in lateral half, extend-
ing to lateral margins of hind coxae by narrow angular tongue, femoral plates
laterally shorter than coxae. Abdomen short, apex rounded, last sternite
distinctly longer than remaining sternites, its basal width approximately
two-thirds of abdominal base. Legs short, femoral apices slightly extending
beyond lateral body margins. Elytra smooth or with numerous rows of large
punctures fusing into longitudinal meandering grooves.

Species composition. Two species in Early Cretaceous of Trans-Baikal.

Comparison. Distinguished from other genera by the shape of femoral
plates whose length is sharply reduced laterally but extending to sides of
coxae; differs from Lithorabus by posteriorly tapering metepisterna.

Cretorabus capitatus Ponomarenko, sp. nov.
(Plate VII, Photo 7; Figure 47)

Species name coined from ‘capitatus’ (Latin)—large headed.

Holotype. No. 3064/862, PIN, impression of a beetle with displaced
sclerites, distal parts of legs and tip of abdomen absent. Trans-Baikal, Buryat
ASSR, Eravnin region, left bank of Vitim river downstream from the mouth
of Baisa river (Baisa site). Lower Cretaceous, Valanginian-Goteriv, Zazin
series.

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113

Material. Holotype.

Description. Small beetle with flattened body. Length of head two-
thirds its width, untapered all the way to anterior margin of eyes. Eyes large,
strongly protruding, longer than temples. Mandibles short, blunt. Gular plate
twice as long as wide, slightly narrowing at middle, much narrower than
submentum, submentum shorter than mentum, lateral lobes of mentum short.
Antennae extending only up to base of pronotum, their segments almost
cylindrical, proximal segments one-and-a-half times longer than their width,
distal segments scarcely longer than wide. First antennal segment narrower
than distal flagellar segments, longer than third; third same as fourth in size;
flagellar segments equal in length; apical segment almost spherical. Width
of pronotum 1.8 times its length, same length as head and only a little wider
than it, slightly narrowing anteriorly in anterior third, constricted in posterior
third, anterior and posterior angles acute, prolonged. Anterior margin of
pronotum very weakly roundly emarginate at middle, at anterior angles with
small, rather deep recess, lateral margin margined, longitudinal grooves
passing through middle of disk. Prosternum in front of forecoxae almost
one-and-a-half times longer than them; prosternal process small, two-fifths
of coxae in width; anteriorly extending on prosternum as flat projection,

|

Fig. 47. Cretorabus capitatus, sp. nov.; holotype PIN No. 3064/862: a—dorsal
view; b—ventral view of head and thorax. Baisa, Lower Cretaceous.

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114

laterally bounded by sharp planes; oblique raised lines proceeding laterally
and posteriorly from the anterior corners of this projection to the pleura.
Propleura wide, slightly narrowing anteriorly, one-and-a-half times longer
than wide. Forecoxae rounded. Mesosternum only slightly shorter than
middle coxae, mesepisterna broad, rhomboidal, mesepimera short, taenioid,
slightly broadening laterally. Middle coxae larger, rounded, contiguous
under united processes and meso- and metasterna. Scutellum triangular.
Length of metasternum less than its width at posterior margin, anterior
margin half the width of posterior; distance between middle and hind coxae
one-and-a-half times large than middle coxae. Posterior margin of metaster-
num projecting from middle backward at acute angle. Length of hind coxae
two-thirds the width, laterally slightly reduced, lateral margin half the
medial, raised part only a little shorter than femoral plate. Length of femoral
plate two-thirds the coxal width, its projection rounded posteriorly, postero-
lateral corner not strongly extending, lateral margin emarginate. Abdomen
one-and-a-half times longer than meso- and metathorax together, sutures
between basal sternites faint, three posterior most sternites shorter than basal
sternites. Femora short and broad, approximately twice as long as wide, hind
femora clavate. Tibiae scarcely broadening toward apex, shorter than femora,
foretibiae with groove and anteriorly broadening apex forming apparatus for
cleaning antennae, hind tibiae preapically with spines and longitudinal band.
Elytra almost smooth with indistinct interweaving longitudinal rows of
punctures. Punctation fine and weak.

Dimensions. Body length 7.3 mm, width 3.1 mm; elytral length4.2 mm.

Comparison. Distinguished by less wide body, longer metasternum and
smaller size.

Cretorabus latus Ponomarenko, sp. nov.
(Plate VII, Photo 8; Figure 48)

Species name coined from ‘latus’ (Latin)—broad.

Holotype. No. 3064/852, PIN, impression of metasternum, abdomen
and elytra of a beeile. Trans-Baikal, Buryat ASSR, Eravnin region, left bank
of Vitim river downstream from the mouth of Baisa (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Rather large beetle. Length of metasternum one-third the
width at posterior margin; distance between middle and hind coxae only.
slightly longer than middle coxae. Posterior margin of metasternum project-
ing backward at acute angle. Mesepisternum wide, slightly broadening in
anterior half, sharply tapering in front of posterior end and with narrow
angular tongue directed posteriorly and mesally toward lateral comers of
hind coxae. Narrow tongue of mesepimeron also reaching this point. Possibly

115

85 the posterior projections of the metapleurites were actually internal and were
not visible externally in the beetle. Width of hind coxae 1.8 times their length,
their length laterally rather strongly reduced, lateral margin one-third the
mesal, projecting part of coxa slightly shorter than femoral plate. Width of
coxa 1.3 times that of femoral plate; its projecting part rounded posteriorly
and laterally, lateral margin almost without emargination. Sutures between
three basal sternites of abdomen noticeable only laterally, last sternite longer
than remaining sternites, its length one-third the width. Elytra with long-
itudinal rows of large punctures. Punctation rather coarse and dense, par-
ticularly on metasternum.

Dimensions. Body length about 15 mm, width 7 mm; elytral length 9.8
mm.
Comparison. Distinguished by broad body and short metastemum.

CARABIDAE INCERTAE SEDIS
Tribe CONJUNCTIINI Ponomarenko, trib. nov.

Diagnosis. Clypeus not extending to antennal socket. Forecoxal cavities
closed. Meso- and metasterna contiguous lateral to middle coxae, mesopleura
not extending to their cavities. Hind coxae not dividing metapleura and
abdomen, metepisterna with mesal process. Both spurs of foretibiae apical.

Composition. Two genera in the Mesozoic of Asia: Mesorabus in the
Late Jurassic of South Kazakhstan, and Conjunctia in the Early Cretaceous
of Trans-Baikal.

Comparison. Distinguished from other beetles by the apical position of
both spurs of foretibiae and closed middle coxal cavities.

Fig. 48. Cretorabus latus, sp. nov.; holotype PIN No. 3064/852. Baisa, Lower
Cretaceous.

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116

Remarks. Inthe fossil material itis very difficult to ascertain the position
of the depression for cleaning the antennae. Both spurs of foretibiae are
apical, but we cannot entirely rule out the possibility that the groove for
cleaning the antennae passes between them as in Cychrus. It is consequently
also impossible to unequivocally state whether the described group belongs
to Isochaeta or Simplicia. Judging from the character of the inner margin of
tibia in Conjunctia and mesepisterna with mesal process, it is more probably
affiliated to Isochaeta. In any case, based on the structure of foretibiae this
group belongs to the most primitive [carabids], while on the basis of the
structure of pro- and mesosterna it can be included among the most advanced
carabids.

Genus Mesorabus Ponomarenko, gen. nov.

Genus name coined from ‘mesos’ (Greek)—middle, and genus Carabus.

Type species. M. elongatus, sp. nov. Upper Jurassic of South Kazakh-
stan.

Description. Rather large, elongate, cylindrical beetles with large
projecting mandibles. Head large, slightly tapering anteriorly. Genae shorter
than temples. Gular plates narrow. Antennae with short monoliform seg-
ments. Mandibles large, asymmetric. Pronotum shorter than head, transverse,
slightly constricted anterior to the base. Propleura longer than wide. Proster-
nal process narrow, apically rounded. Mesosternum longer than middle
coxae, metasternum rather long, slightly tapering anteriorly, its posterior
margin almost straight, transverse suture indistinct. Abdomen rather long.
Legs short, spurs of foretibiae apical, three middle segments of hind tarsi
short and broad.

Species composition. Monotypic genus.

Comparison. Differs from the genus Conjunctia in long mandibles,
bluntly rounded apex of prosternal process, almost straight posterior margin
of metasternum and short middle segments of hind tarsi.

Mesorabus elongatus Ponomarenko, sp. nov.
(Plate VIII, Photo 1; Figure 49a, b)

Species name coined from ‘elongatus’ (Latin)—elongated.

Holotype. No. 2784/1525, PIN, impression of a beetle without large part
of legs. South Kazakhstan, Chimkent oblast, Algabass region, south-western
flank of Kashkarata river valley, Aulie area near Mikhailovka village |
(Karatau-Mikhailovka site). Upper Jurassic, Karastau* series.

Material. Holotype.

*So given in the Russian original. Based on earlier references to this series it should read
Karabastau series—General Editor.

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117

Description. Maximum width of body beyond shoulders. Head capsule
equal in length and width, tapering in front of eyes. Genae half the length of
eyes. Labrum anteriorly emarginate. Mandibles much longer than wide,
apically curved. Submental lobes rather broad, notch between them shallow.
Antennae scarcely extending beyond pronotal base, their first segment broad
and short; second much narrower than it, almost equal in length and width;
third equal to fourth and fifth together; all segments from fourth onward
almost equal in length, somewhat broadening apicaily. Pronotum equal in
length to head capsule, its lateral length two-thirds the maximum width;
anterior margin of pronotum with shallow straight emargination. Propleura
twice as long as wide, prosternum in front of forecoxae noticeably shorter
than them. Prosternal intercoxal process much narrower than coxae, some-
what broadening beyond them, apically rounded. Forecoxae angular.
Mesosternum longer than middle coxae, mesepisterna almost square,
mesepimera half the length of middle coxae, almost not ionger than them.
Length of metasternum half the width at posterior margin, anterior margin
two-thirds the width at posterior, distance between coxae* 1.3 times longer
than middle coxae, almost one-third the width at posterior margin of
metasternum. Metepisterna slightly broadening anteriorly, posteriorly taper-
ing strongly. Mesal processes of metepimera rather wide, extending to lateral
corners of hind coxae. Hind coxae triangular, length 1.7 times the width; part
of coxae projecting above abdominal plane longer than their width, posterior-
ly rounded, laterally emarginate. Abdomen almost two times longer than
meso- and metathorax together, tapering from base of fourth visible sternite,
third sternite longest, remaining sternites shorter, sutures between sternites
straight. Legs short, femora uniformly thickened, length of hind femora
three-times their width. Forefemora and tibiae equal in length, short, hind
femora markedly longer than forefemora, shorter than tibiae, tibiae gradually
broadening toward apex. Spurs of hind tibiae slightly shorter than first tarsal
segment. Tibiae 1.3 times the length of hind tarsi, first tarsal segment equal
to fifth or second and third together; second to fourth equal, noticeably
broader than remaining segments. Integument with fine punctation, almost
smooth, punctures larger only on elytral disk.

Dimensions. Body length 21 mm, width 8 mm; elytral length 12.5 mm.

Genus Conjunctia Ponomarenko, gen. nov.

Genus name coined from ‘conjunctia’ (Latin)—connection.
Type species. C. prodroma, sp. nov. Lower Cretaceous of Trans-Baikal.

*So given in the Russian original. By context, this would imply only hind coxae, but its
comparison with middle coxae raises doubt. By analogy with earlier descriptions, it is more likely
that the reference is made to the distance between the middle and hind coxae—General Editor.

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118

Description. Small, elongate, cylindrical beetle, head large, elongate,
with projecting mandibles, genae much shorter than temples. Gular plate
narrow. Antennae with short weakly moniliform segments. Pronotum shorter
than head, transverse, weakly constricted in front of base. Propleura longer
than wide. Prosternal process narrow. Mesosternum long, posteriorly an-
gularly extending, its transverse suture indistinct. Hind coxae oblique, short,
slightly separated, with weakly developed raised part. Abdomen broad and
short, terminal sternites telescopic. Legs rather short, femora thickened,
tibiae slightly broadening toward apex, both spurs of foretibiae apical, at
same level, inner side of tibiae with groove for cleaning antennae. Segments
of fore- and middle tarsi short, not broadening.

Species composition. Monotypic genus. |

Comparison. Distinguished by narrow prosternal process, posteriorly
angularly extending hind margin of metasternum, and a short abdomen.

Fig. 49. Representatives of Conjunctionini*: a, b —Mesorabus elongatus, sp. nov.;

holotype PIN No. 2784/1525: a—dorsal view; b—ventral view; Karatau, Upper

Jurassic; c—Conjunctia prodroma sp. nov.; holotype PIN No. 1668/1763. Baisa,
Lower Cretaceous.

*Tribe name differs from what has been given on pagel 15—General Editor.

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119

Conjunctia prodroma Ponomarenko, sp. nov.
(Plate VIII, Photo 2; Figure 49c)

Species name coined from ‘prodroma’ (Greek)—running ahead.

Holotype. No. 1668/1763, PIN, impression of an almost entire beetle.
Trans-Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river
downstream from the mouth of Baisa river (Baisa site). Lower Cretaceous,
Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Beetle with flattened, posteriorly broadening body with
maximum width near hind coxae. Head scarcely shorter than its width,
slightly broadening beyond eyes, genae very short, eyes slightly longer than
temples. Antennae attached near anterior margin of eyes. Antennal segments
short, first segment twice as thick as remaining segments, second differing
slightly in size from distal segments. Gular plate narrowing at middle, here
its width almost half the base and one-third its maximum width. Submental
lobes narrow, incision between them shallow. Pronotum two-thirds the
length of head, its length almost half the width, width in front equal to head,
with transverse line in front of posterior corners, anterior margin wider than
posterior. Propleura more than twice as long as wide, prosternum in front of
coxae one-and-a-half times longer than them. Prosternal process much
narrower than coxae, anierior to coxae rounded. Mesosternum one-and-a-
half times longer than middie coxae, mesepisterna almost square,
mesepimera short, taenioid. Distance between middle coxae and pleura not
less than the width of intercoxal process. Metasternum half the width at
posterior margin, width of anterior margin half of posterior; distance between
hind coxae half their length. Metepisterna narrow, anteriorly sharply
broadening, posteriorly proceeding as pointed narrow extension to lateral
comers of hind coxae, mesal process of metepimera narrow. Width of hind
coxae 3.5 times their length, raised part of coxa very slightly shorter than its
width, laterally strongly emarginate. Abdomen approximately equal in length
to meso- and metathorax together, five basal sternites equal in length, last
sternite two-and-a-half times longer, its anterior margin projecting. Apex of
abdomen blunt, rounded. All femora almost equal in length. Foretibiae equal
to femora in length, slightly broadening apically, at apex with an internal
process, and emarginate more proximally in apical third. Basal segment of
foretarsi longer than the two following. Middle tibiae shorter than femora,
scarcely broadening toward apex. Middle tarsi shorter than tibiae, first
segment equal to second and third together. Hind tibiae slender, noticeably
shorter than femora, basal tarsal segments long, slender. Integument with
sparse, fine punctation, almost glabrous.

Dimensions. Body length 5.1 mm, width 2.1 mm; elytral length 3.1 mm.

89

88

120
Formal genus “Carabites”

“Carabites” vitimensis Ponomarenko, sp. nov.
(Plate VIII, Photo 3; Figure 50a)

Holotype. No. 3064/871, PIN, impression of a beetle without head,
prothorax and legs. Trans-Baikal, Buryat ASSR, Eravnin region, left bank
of Vitim river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Small, flat, broad beetle. Mesosternum not longer than
middle coxae, with depression for the apex of prosternal process. Mesepister-
na almost square, only slightly short of reaching middle coxal cavities,
mesepimera extending up to middle coxal cavities, rather long, laterally a

Z eae / y)
Se \ IC |

Fig. 50. Carabidae incertae sedis: a—“Carabites” vitimensis, sp. nov.; holotype
PIN No. 3064/871; Baisa, Lower Cretaceous; b, c—‘Carabites” creta, sp. nov.;
holotype PIN No. 2383/111; Kzyl-Dzhar, Upper Cretaceous; d—Harpalinae,
gen. sp.; specimen PIN No. 2383/206a, foretibia; Kzyl-Dzhar, Upper Cretaceous.

== =

121

little longer. Anterior projection* broad, extending to anterior margins of
middle coxae. Middle coxae set apart, transverse, oval. Length of metaster-
num approximately one-third its width at posterior margin, anterior margin
half the width of posterior. Distance between [middle and hind] coxae
one-and-a-half times the length [of middle coxae]. Metepisternum triangular,
its inner margin almost straight, width at anterior margin half the length,
posterior margin very narrow. Hind coxae triangular, contiguous, their
maximum length one-third the width. Processes of femoral plates absent.
Abdomen short and broad, its apex blunt, terminal sternites articulate,
telescoping. Elytra with a few grooves, bearing large punctures. Integument
almost smooth.

Dimensions. Body length 7 mm, width 3.5 mm; elytral length 4.3 mm.

Taxonomic position. The impression described here is the fossil remnant
of a beetle, all characters of which correspond to those of the present-day
carabids. However, its imperfect preservation does not allow formal com-
parison with the present-day forms.

“Carabites” creta Ponomarenko, sp. nov.
(Plate VIII, Photo 4; Figure 50b, c)

Holotype. No. 2383/111, PIN, isolated sclerites of a beetle, head with
pronotum, meso- and metasterna, two (middle) ? legs. South Kazakhstan,
Kyzyl-Ordyn oblast, Chiilii region, north-western spurs of Karatau mountain
range (Kyzyl-Dzhar site). Upper Cretaceous, Turonian, Beleutin series.

Material. Holotype.

Description. Small beetle with long narrow head and long cordate
pronotum. Length of head capsule slightly more than its maximum width at
the level of eyes, tapering from eyes anteriorly and posteriorly. Genae shorter
than eyes, temples longer than them. Sharp keel proceeding from sides of
head over the eyes to the base of antennae. Basal antennal segment long and
slender. Length of pronotum very slightly shorter than width, rounded
anteriorly and with shallow emargination, rather strongly constricted in front
of posterior angles, posterior margin straight, maximum width of pronotum
anterior to middle. Meso- and metathorax together shorter than prothorax.
Middle coxae set apart, mesepisterna of almost same length and breadth,
mesepimera rather long, laterally broadening, extending to middle coxal
cavities, middle coxae set apart**. Length of metasternum half the width at
posterior margin, its anterior margin half the width at posterior, metasternal
suture straight, posterior margin of metasternum angularly extending back-
ward. Metepisternum strongly broadening anteriorly, its width posteriorly

*Of mesepimeron—General Editor.
** An obvious repetition in the Russian original—General Editor.

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122

three fourth the length. Metepimeron visible posteriorly between metepister-
num and abdomen as a small rectangular sclerite. Legs rather short, femora
only slightly extending beyond lateral body margins. Femora uniformly
thickened, approximately four times longer than wide. Tibiae almost linear,
most frequently flat, a little shorter than femora, with longitudinal ridge or
groove, spurs of middle tibiae small. Tibiae apically with numerous small
spines, large spines absent. Middle tarsi slender, equal to tibiae in length,
their first segment longest, third and fourth segments shortest. Tarsal seg-
ments densely covered with small spines. Head and pronotum smooth,
metasternum with large, dense tubercles.

Dimensions. Body length nearly 10 mm, length of head and pronotum
3.6 mm, length of meso- and metasterna together approximately 3 mm.

Taxonomic position. Due to imperfect preservation of the beetle it is not
possible to precisely establish its taxonomic position. Nevertheless, it merits
description as one of the earliest advanced carabids of the present-day type.
It strongly resembles specialized present-day mollusk-eaters in the shape of
its pronotum and head. Judging from the open middle coxae, exposed
metepimeron, short legs and non-fossorial tibiae it must be affiliated to
Isochaeta but this cannot be confirmed due to the lack of possibility of
studying the apical structure of foretibiae and prosternum. Moreover, it is not
definite that the small sclerite at the junction of metepisternum and hind coxa
is actually the mesal part of the metepimeron.

Harpalinae, gen. sp.
(Plate VIII, Photo 5; Figure 50d)

Specimen No. 2383/206a, PIN, foretibia. South Kazakhstan, Kyzyl-Ordyn
oblast, Chiilii region, north-western spurs of Karatau mountain range (Kyzyl-
Dzhar site). Upper Cretaceous, Turonian, Beleutin series.

Description. Isolated foretibia of ground beetle, without apex. Small
emargination in distal part of tibia, at proximal end [of emargination] large
spur inserted, blocking it. Apex of spur not preserved. Two rows of large
fossae for spine articulation along tibia; proximally with three grooves for
cleaning antennae, and six along the opposite margin of tibia.

Taxonomic position. Judging from the nature of the groove for cleaning
the antennae, the tibia belongs to a beetle of subfamily Harpalinae (sensu
Crowson, 1955). This is the most ancient find of Harpalinae.

ADEPHAGA INCERTAE SEDIS

Genus Necronectulus Ponomarenko, gen. nov.

Genus name coined from genus Necronectes.

9

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123

Type species. N. avus, sp. nov. Lower Jurassic of East Kazakhstan.

Description. Small, oval beetle. Head transverse, markedly longer than
pronotum. Eyes rather large, longer than temples. Gular plate almost equal
in length and width, somewhat narrower at middle. Antennae slender and
short, extending only to base of pronotum, their segments cylindrical.
Pronotum anteriorly slightly emarginate, much shorter than its width,
anteriorly roundly tapering, its base not narrower than elytra at shoulders.
Propleura almost equal in length and width. Prosternal process noticeably
projecting beyond forecoxae. Forecoxal cavities open. Mesosternum short,
with medial triangular depression for prosternal process. Mesepimera and
mesepisterna almost equal. Metasternum transverse, strongly roundly taper-
ing anteriorly. Middle coxae set apart. Metepisterna strongly broadening
anteriorly, extending to midcoxal cavities. Transverse metasternal suture
distinct, angularly extending backward. Hind coxae oblique, laterally com-
pletely separating metapleura and abdomen; large femoral plates present.
Femoral plates transverse, laterally gradually tapered. Abdomen long, ar-
ticulation of its sternites free. Elytra without grooves, with interweaving rows
of large punctures.

Species composition. Monotypic genus.

Taxonomic position. Formally, judging from the metepisterna extend-
ing to middle coxa! cavities and the structure of hind coxae, it must be related
to subfamily Eodromeinae of the family Trachypacheidae. However, all the
representatives of the latter are terrestrial whereas the fossil described here
is more that of an aquatic beetle. The general body shape and slender filiform
antennae are indicative of this. The short propleura are also more charac-
teristic of aquatic beetles. Unfortunately, the structure of the legs of
Necronectulus is not known, hence it is not possible to determine its precise
taxonomic position. It is noteworthy that among the present-day carabids,
the representatives of Pseudomorphini morphologically resemble aquatic
beetles no less than Necronectulus aithough they also lead a terrestrial mode
of life.

Necronectulus avus Ponomarenko, sp. nov.
(Plate VIII, Photo 6; Figure 51)

Species name coined from ‘avus’ (Latin)—ancestor.

Holotype. No. 1362/27, PIN, impression of a beetle without legs. East-
Kazakhstan oblast, Zaisan region, Saur mountain range, right bank of Ak-
kolka river, tributary of Karaungir (Kenderlyk site). Lower Jurassic, Tologoi
series.

Material. Holotype.

Description. Length of head two-thirds its basal width, tapering in front
of eyes, labrum truncate. Genae almost equal to temples in length. Mandibles

92

91

124

rather long, acute. Lateral lobes of submentum short, rounded. First antennal
segment slightly thicker than the remaining segments, equal to last two
together; third segment not longer than fourth; length of flagellar segments
approximately three times the width. Pronotum slightly overlapping the head,
anteriorly with shallow rounded emargination. Prosternal length anterior to
forecoxae and the length of coxae equal. Prosternal process slightly narrower
than coxae, broadening beyond their middle, apically rounded. Propleura
posteriorly very slightly broader than long, provided with pointed mesal
process, partially covering forecoxal cavities. Mesosternum in front of
middle coxae not shorter than them. Length of metasternum one-third the
width at posterior margin, anterior margin less than half the width of
posterior. Distance between middle and hind coxae one-and-a-half times
greater than the length of middle coxae. Metepisterna gradually broadening
in anterior half, their width anteriorly two-thirds the length. Hind coxae
together with femoral plates in length two-thirds the width, reduced in lateral
half, laterally short, taenioid. Abdomen tapering almost from base, last
abdominal sternite at base half the width of abdominal base, its length
two-fifths the width. Apex of abdomen blunt. Body with fine punctation
dorsally. Elytra with shallow punctures not forming regular rows. Ventral

a b

Fig. 51. Necronectulus avus, sp. nov.; holotype PIN No. 1361/27*; a—dorsal
view; b—vental view. Kenderlyk, Lower Jurassic.

*Given as 1362/27 in the Text and Plate VII1I—General Editor.

125

side with deeper punctures, metasternum with large punctures, fusing into
transverse grooves. Thoracic pleura, femoral plates and anterior halves of
abdominal sternites with smaller and sparser punctures, posterior halves of
abdominal sternites with very small punctures, almost smooth.
Dimensions. Body length 4.2 mm, width 2.0 mm, elytral length 3.0 mm.

Genus Cretotaenia Ponomarenko, gen. nov.

Genus name coined from ‘creta’ (Latin)—chalk and ‘taenia’ (Greek)—band.
Type species. C. pallida, sp. nov. Lower Cretaceous. Trans-Baikal.
Description. Small, elongate aquatic larvae with ambulatory legs and

rather long urogomphi. Head almost square, slightly tapering posteriorly,

noticeably shorter dorsally than ventrally, hence occipital foramen shifted to
dorsal side, while head was directed forward and upward during life. Parictal
suture short, temporal sutures commencing almost from posterior margin of
head. Ocular sclerites* large, occupying almost one-third the length of head
capsule, with six large, close-set lateral ocelli. Anterior margin of head
capsule without nasale, with large shallow emargination. Antennae shorter
than mandibles and head capsule, their segments thick. Mandibles slender
with pointed curved apices, retinaculum rather large, positioned close to the
midlength of mandible. Maxillary palp four-segmented, galea two-seg-
mented. Labium short, transverse, anteriorly projecting, without distinct
ligula, its palp two-segmented. Across the head capsule there is highly
sclerotized area in the form of an inverted “T”—apparently the hypopharynx.

Thoracic segments almost equal, weakly sclerotized, especially the
prothorax. Mesothoracic spiracles slightly larger than abdominal spiracles,
metathoracic spiracles very small, inconspicuous. Legs with rather long,
cylindrical, weakly sclerotized segments.

Abdominal segments with small rounded or oval, isolated sclerotized
patches, two larger on each tergite and two smaller on pleura or sternites close
to spiracles. Spiracles of seven anterior abdominal segments small, that of
the last segment (at posterior margin of eighth) much larger than them,
converging at a distance not exceeding diameter of sclerotized patch on
tergite. Ninth tergite sclerotized, pygopods short, urogomphi long, distinctly
segmented.

Species composition. Monotypic genus.

Taxonomic position. Based on body segmentation, Cretotaenia is most
closely related to carabids. However, this cannot be formally proved because
of the lack of an opportunity to study the structure of the legs in detail.
Moreover, aquatic metapneustic forms are absent among the larvae of
carabids while the sclerotized patches on the tergites, represented by small
round spots, are observed only in larvae of tiger beetles.

*See our note on page 25—General Editor.

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126

Cretotaenia differs from all air-breathing larvae of other aquatic
Adephaga by the presence of a distinct tergite on the ninth abdominal
segment. Among Polyphaga, air-breathing aquatic larvae occur in Hydro-
philidae but lack such long segmented urogomphi and the posterior spiracles
are differently constructed. The externally similar larvae of Staphylinidae are
not aquatic and lack such a large retinaculum on the mandibles. Nevertheless,
this larva is most probably the larva of a beetle of the suborder Adephaga,
combining the characters of both Geadephaga and Hydradephaga.

Cretotaenia pallida Ponomarenko, sp. nov.
(Plate VIII, Photos 7-9; Figure 52)

Species name coined from ‘pallida’ (Latin)—pale.

Holotype. No. 1989/2890, PIN, entire late-instar larva. Trans-Baikal,
Buryat ASSR, Eravnin region, left bank of Vitim river downstream from the
mouth of Baisa river. Lower Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Thirty larvae from a single site, mostly entire, but due to the
weak sclerotization the structural details could be examined in only a few
impressions. All impressions were gathered from a single layer (Martinson,
1961).

Description. Small or minute larvae with very thin integument so that
in the impressions only the following were usually visible: sclerites* of
lateral ocelli, sclerotized hypopharynx, sclerotized patches on occiput and
anterior margins of meso- and metathoracic tergites and round, paired
sclerotized patches on abdominal segments, and main longitudinal tracheal
trunks (Fig. 52a, b). Head (Fig. 52c) noticeably tapering posteriorly, dorsal
length two-thirds of ventral. Coronal suture three-fourths the length of head
capsule dorsally, portion of head capsule between frontal sutures triangular.
Dimensions of lateral ocelli exceeding the distance between them.
Sclerotized patches on parietals, sites of attachment of massator muscles
clearly distinguishable on the weakly sclerotized head capsule. First antennal
segment transverse, second longest, longer than third and fourth, length of
third almost twice fourth. Length of mandible not less than head capsule
dorsally, longer than antenna, width of mandible proximal to retinaculum
one-third its length, width distal to retinaculum half of proximal, curved
inward, with pointed, elongated apices. Maxilla together with palp equal in
length to mandible, stipes slightly narrower than proximal part of mandible.

Prothorax noticeably shorter than head, anteriorly tapering, meso- and
metathorax slightly shorter than it. Femora and tibiae of all legs approximate-
ly equal in length, tarsi somewhat shorter. Judging from the position of
sclerotized patches, all abdominal segments except the ninth almost equal in
length, seventh and eighth narrower than preceding segments, ninth half the

*See note on page 25—General Editor.

95

127

length of eighth. Urogomphi 5 times longer than ninth abdominal segment.
Sclerotized patches on abdominal tergites oval in late-instar larvae, elongate
longitudinally, almost round in early-instar larvae. Sclerotized patches on
pleurites rounded, not exceeding size of spiracle peritremes. Main lon-
gitudinal tracheal trunks thick, slightly narrower than the peritremes of the
eighth abdominal segment and much wider than the peritremes of the
remaining spiracles; thickness of remaining tracheae greatly exceeding them.
Perimeters of spiracles highly sclerotized.

Dimensions. The larvae could be divided into two size classes. The
smaller larvae differ from the larger in relatively large head, urogomphi, and
sclerotized patches of cuticle. This is usually characteristic of early-instar
larvae. It is therefore convenient to consider this material as composed of
late- and early-instar larvae of a single species of beetle and not as belonging
to different species:

Stage Number Length without Width of Diameter of
urogomphi, head, mm sclerotised
mm patches
on tergites,
mm
Late 19 av. 12.3 av. 1.0 av. 0.43
8.0-14.3 0.8-1.1 0.4-0.5
Early 11 av. 6.1 av. 0.45 av. 0.18
4.7-7.0 0.5-0.6 0.17—0.20

Ecology. Posterior spiracles of larvae significantly larger than remain-
ing spiracles and shifted to postero-dorsal margin of tergite which, along with
the highly sclerotized very large tracheae, indicates that these were aquatic
air-breathing larvae. At the same time, the same degree of sclerotization of
all the peritremes, and rather large tracheal trunks extending to all the
spiracles, suggests that possibly all the spiracles were functional and the
larvae could live on land at least for a short time. Judging from the very weak
sclerotization, the larvae could exist only in moist conditions, for example,
in soil. These larvae were undoubtedly predatory. Some device for injecting
digestive juices into the prey, a characteristic feature of most aquatic
Adephaga, is absent in them, and during feeding they possibly held the prey
above the water surface as hydrophilids do. The uplifted position of the head
is also possibly associated with this since it is also characteristic of
hydrophilids. No special natatory device is visible in the larvae, possibly
because the weak sclerotization of the legs prevented their detailed study.
However, the low level of sclerotization itself may indicate their weakness
and poor locomotory activity.

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94 Fig. 52. Cretotaenia pallida, sp. nov.: a—holotype PIN No. 1989/2890, head.
Paratypes: b—PIN No. 1989-2889, c—PIN No. 1989/2903, d—PIN No.
1989/2893, head and thorax, e—PIN No. 1668/1837, the same, f—PIN No. 1989/
2892, tip of abdomen, g—PIN No. 1989/2900, the same. Baisa, lower Cretaceous.

95;

129

Among the adult aquatic beetles found in this site, none could be
definitely considered Cretotaenia. These were probably similar to terrestrial
beetles like Eodromeus (family Trachypacheidae) in which many hydra-
dephagan features are characteristically present. These beetles are very
frequently found buried and most probably led a littoral mode of life.

Genus Triadogyrus Ponomarenko, gen. nov.

Genus name coined from Triassic, and ‘gyrus’ (Greek)—spin.

Type species. T. sternalis, sp. nov. Triassic of Soviet Central Asia.

Description. Mesosternum noticeably longer than middle coxae. Middle
coxae equal in length and width; distance between middle and hind coxae
equal to the length of middle coxae, metepisterna not extending to middle
coxal cavities. Hind coxae short, without femoral plates. Abdomen with six
visible sternites. Eighth sternite divided into two halves, concealed. Elytra
without punctate furrows.

Species composition. Monotypic genus.

Taxonomic position. Distinctly differing from all Early-Mesozoic Ade-
phaga in its advanced form so that formal characterization cannot completely
affiliate it even to ground beetles. Also, this beetle was almost undoubtedly
aquatic. This is indicated by the characteristic structure of its integument,
body shape, and very large spiracles, particularly the latter. Mesosternum
long, longer than practically in any other Mesozoic representative of

Fig. 53. Triadogyrus sternalis, sp. nov.; holotype PIN No. 3320/13. Garazhovka,
Upper Triassic.

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130

Adephaga, except whirligigs; structure of hind coxae resembling that of the
most primitive whirligigs. This form is probably closely related to the
ancestors of whirligigs.

Triadogyrus sternalis Ponomarenko, sp. nov.
(Plate III, Photo 9; Figure 53)

Species name coined from ‘sternum’ (Latin)—chest.

Holotype. No. 3320/13, PIN, impression of beetle without head,
prothorax or legs. Ukraine SSR, Khar’kov oblast, Izyum region, mouth of
Bereni river near Garazhovka settlement. Upper Triassic, Protopopov series.

Material. Holotype.

Description. Length of mesosternum half the width; mesopleura large,
almost square; mesepisterna much larger than mesepimera. Length of
metasternum one-third its width, posterior margin acutely angularly project-
ing backward. Length of hind coxae half their width. Spiracles one-fourth
the length of sternites, last spiracle twice as large as preceding spiracles.

Dimensions. Body length approximately 13 mm, width 5.6 mm; elytral
length 10.2 mm.

MESOZOIC STAGES IN THE EVOLUTION
OF ADEPHAGA

The evolution of beetles of the suborder Adephaga differs radically from the
chronoclinal changes in other suborders of beetles. Beetles of the suborder
Archostemata developed as a single, rather compact group in the Permian
and Mesozoic, preserving the same trends even in the relictual groups which
have survived to the present. Of course, well-defined stages can be distin-
guished in this single process, but general morphological as well as ecologi-
cal trends are preserved independently. Individual phylogenetic branches can
be traced throughout the entire period of existence of the group. Therefore it
is impossible to speak of temporal associations of the Archostemata as some
stage in the development of this group as a whole. A similar situation is seen
in the Polyphaga, with the only difference being that they evolved during the
Mesozoic and Cenozoic eras. It can be said that extensive development of
higher beetles (Cucujiformia) is more characteristic of the Cenozoic, whereas
ihe less advanced forms were primarily distributed in the Mesozoic. How-
ever, all the main lineages of the Polyphaga were already widely represented
in the Mesozoic, and the tendencies within these groups have been preserved
without change.

The evolution of the Adephaga is totally different. Here the Mesozoic
forms constitute a sufficiently unified morphological stage, although their

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131

developmentalso proceeds along many branches. The Mesozoic fauna of the
Adephaga consists almost exclusively of representatives of dead-end strictly
Mesozcic branches, which became extinct toward the end of the Mesophytic
(i.e., toward the middle of the Cretaceous), as did most other strictly
Mesozoic groups of terrestrial animals. In the Late Cretaceous we suddenly
begin to observe representatives of the essentially modern, Cenozoic fauna.
Among the earliest (Triassic) Adephaga, the same forms can be observed as
in the Cretaceous, though in smali numbers. This certainly does not suggest
that the Adephaga did not evolve in the Mesozoic. On the contrary, their
evolution in individual structural features, primarily the locomotory organs,
can be traced fairly well. Among the Mesozoic Adephaga, beetles such as
Coptoclava were highly specialized for an aquatic mode of life
(Ponomarenko, 1975). However, this far reaching specialization occurred in
them at a primitive level of general organization.

For beetles of the suborder Adephaga, and more significantly for all
Oligoneoptera, the mode of life of the larva and adult is similar, and
consequently the evolutionary trends are identical for both life stages. In view
of this, the Adephaga have highly prothetelic larvae. These are relatively
free-living, usually active predators with fully segmented long legs, and
usually compact, highly sclerotized integument. This larval type is conven-
tionally considered to be primarily ancestral, although this assumption may
be false. Unfortunately, paleontological material on the larvae of holo-
metabolous insects is very rare, and larvae are known for only a few,
primarily aquatic, species. However, this may be viewed as evidence in favor
of the secondary prothetely of adephagan larvae. Cretotaenia is an aquatic
larva with an indeterminate taxonomic position, having rather thin integu-
ment with small plate-like sclerotized patches. A similar structure is observed
only in the larvae of tiger beetles. The latter constitute a group in which
features of distinct specialization combine with a common, rather low level
of organization. It can therefore be assumed that this similarity is not
accidental but reflects at least one of the modes of larval prothetely in the
Adephaga. Cretotaenia and tiger beetles were hardly closely related and this
trend is found fairly commonly in the suborder, and seen even in its distant
branches.

At present, the structure of the aquatic larvae is best known among the
larvae of the Mesozoic Adephaga. In fact, they are all related to the dytiscoid
branch of the Hydradephaga. Larvae of the Mesozoic Gyrinidae and the
Mesozoic ancestors of the Haliplidae have not yet been found. Larvae of the
Mesozoic dytiscoids were already highly specialized nektonic (Para-
hygrobia, Stygeonectes, Coptoclava) or benthic (Angaragabus) predators
with telescopic abdominal segments and with aerial respiration through the
spiracles of the last abdominal segment only. Since the Cretotaenia too
represent a distinctly aberrant off-shoot in evolution, the earlier stages of

132

larval evolution in the Adephaga remain completely unknown. The larvae of
the schizophoroid Archostemata, the direct ancestors of the Adephaga, are
also unknown. Hence paleontology does not offer any new clues to the
evolution of the Adephaga larvae, except for the above mentioned suggestion
of a rather low level of sclerotization in the early Adephaga.

We have an incomparably large volume of material at our disposal for
reconstructing the evolution of adult Adephaga, including its early stages.
The origin of the Adephaga from the schizophoroid Archostemata was
postulated earlier (Ponomarenko, 1969b, 1973). This assumption was based
almost exclusively on the study of Archostemata and the present-day
Adephaga. They study of Early Mesozoic Adephaga generally confirmed this
assumption but many details differed from those postulated earlier.

Among the structural features available for paleontological study, only
two characters separate the Adephaga from the Archostemata. Unfortunate-
ly, even based on these characters the suborders are not fully distinguishable
even in present-day material, not to mention fossil remnants. The demarca-
tion between the suborders is very indistinct. Both differentiating features
characterize stages of consolidation of the integument for more rigid com-
binations of sclerites. The first feature is the character of closure of the middle
coxal cavities. In all the known Archostemata the metepisterna extend to the
middle coxal cavities and contribute to the closure of their lateral walls.
Among the present-day Adephaga this type of structure is observed only in
some diving beetles, exceptionally well adapted to an aquatic mode of life
and having legs distinctly specialized for swimming, i.e., in clearly derived
forms. In the lower Adephaga the metepisterna never extend to the middle
coxal cavities.

Among the Archostemata for which an aquatic mode of life can be
postulated, forms exceptionally well adapted for swimming are absent. On
this basis, forms with an external appearance characteristic of terrestrial
beetles, and with metepisterna extending to the middle coxal cavities, were
described as representatives of the Archostemata (Ponomarenko, 1969b).
The study of the Mesozoic Adephaga showed, however, that numerous
terrestrial beetles are present among them, in which the metepisterna extend
to the middie coxal cavities. Moreover, most Mesozoic terrestrial beetles had
this type of structure, including those beetles in which the combination of
characters categorize them as primitive ground beetles. Primitive aquatic
Adephaga with a similar structure to the middle coxal cavities have also been
found. This type of situation was quite expected. If the rather highly advanced
Adephaga preserved the structure of middle coxal cavities characteristic of
ancestral Archostemata, it would be natural to expect that a similar structure
is found in more primitive extinct forms.

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133

For the second feature—viz., the nature of articulation of the metaster-
num, hind coxae and abdomen—an a priori much higher discriminatory
value may be assumed. In the Adephaga, unlike all other beetles, the hind
coxae fully separate the metasternum and abdomen since they are all ar-
ranged in one plane. This condition is undoubtedly derived from the more
widespread one wherein the hind coxae lie above the basal, morphologically
second, abdominal sternite articulating with the metasternum and fully or
partially covering the basal sternite. If under these circumstances the hind
coxae are strongly reduced, then the articulation of the metasternum with the
abdomen, both between the coxae and along their sides, ought to proceed
from the morphologically third abdominal! sternite, and not the second as in
the Adephaga. Several hypotheses have been advanced on the adaptive
significance of the position of hind coxae characteristic of the Adephaga:
adaptation for fast running, for life under bark (Crowson, 1955), and for
swimming (Ponomarenko, 1973a). Functionally, however, these three trends
are very similar. In each, the change is associated with the shift of the femur
as far forward as possible in order to lengthen the stride of the leg in a single
step.

A new difficulty arises during a study of the fossil material. It is
frequently very difficult to accurately interpret the structure of abdominal
base on an impression and determine whether the coxae are superimposed
on the base of the abodmen or accommodated in the cavities in it. Moreover,
before fossilization the body of the beetle would frequently be inflated with
gases arising during putrefaction and the abdomen would be displaced from
under the coxae, imitating the structure characteristic of the Adephaga. Thus
the only method of understanding the interrelations of the coxae and ab-
domen is to carefully study the nature of the surface of the basal abdominal
sternites. The parts of the abdominal sternites hidden by the coxae are almost
always strongly differentiated from the other sternites by their degree of
sclerotizaiion and nature of their punctation. If we observe on the first visible
abdominal sternite on each side of the coxae the same very coarse punctation
that is typical of many Mesozoic beetles, we can consider that this sternite
was not hidden under the coxae and that the abdominal structure is typical
of the Adephaga.

Morphological studies of the abdominal base in Haliplidae are of par-
ticular significance in proving any concepts regarding its characteristic
formation in the Adephaga. In these beetles, the hind coxae with their large
femoral plates apparently restricted the strong forward movement of the
femora. As far as we know, there are no convincing adaptive interpretations
of this extravagant structure. Such a structural peculiarity has no great
bearing on the hypothesis that the hind coxae developed in the Adephaga in
response to life on land, since similar femoral plates could have arisen as a
consequence of a transition to life in water. The meaning of this structural

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feature is far more essential to the concept of an aquatic origin of the
Adephaga and io the structural change in coxae as an adaptation to swim-
ming. In many structural features the Haliplidae are the most primitive
Adephaga. Their structure oughi to be close to the original since a large
number of Adephaga, particularly the Early Mesozoic ones, had large
femoral plates, though notas large as in the Haliplidae. Femoral plates totally
preclude the possibility of a forward extension of the femora, therefore the
idea of inclusion of the lateral parts of hind coxae in the body wall for
promoting this type of movement cannot be advanced. In fact, in the
Haliplidae and Triassic Triaplidae the hind coxae are markedly different from
those in the remaining Adephaga. The hind coxae of Haliplus are obliquely
arranged and the place of attachment of the trochanter is strongly shifted
posteriorly compared to the iateral part of the coxa (Fig. 54). The hind coxae
in Triaplidae have a similar structure. Here neither the femoral plates nor the
lateral part of the coxa obstruct the strong forward extension of the femur.
Correspondingly, even the articulation of the abdomen itself strongly differs
from that in other Adephaga. The second true abdominal sternite is almost
entirely concealed under the hind coxae. Here its structure differs sharply
from that of the other sternites. Only the small triangular areas near the
postero-lateral corners have the same structure as the third sternite. A high
sclerotized ridge runs along the posterior margin of this area and extends
towards the body axis, closely joined to the posterior surface of the hind
coxae. The ridges from both sides of the abdomen converge in the mid-line
into a flat, triangular, posteriorly tapering area with sharply defined margins.
This area proceeds through the third true sternite and extends into the fourth.
The third sternite, divided by the area into two isolated parts, seems to be
identical to the second sternite of the remaining Adephaga. However, it
appears that no one has considered the basic differences in the abdominal
structure of the Haliplidae and the remaining Adephaga. It is noteworthy that
in several other structural features the Haliplidae may be distinguished from
the remaining Adephaga. The apical spiral twisting of the hind wings, which
is characteristic of the Archostemata, is preserved only in them; their larvae
have a unique respiratory apparatus—tracheal microgills (Seeger, 1971).

The Triassic Triaplidae which possess the largest femoral plates after
the Haliplidae, apparently also resemble the latter in the structure of the
abdominal base. Here also a small sclerotized area is seen in the posterolateral
comers of the second true abdominal sternite; this area is structurally similar
to the remaining sternites, however, the triangular area on the third sternite
is absent. The medial part of the hind coxae is markedly prolonged posterior-
ly, the length of the coxa is comparable with its width at the anterior margin.
Therefore, despite the presence of large femoral plates which are almost not
at all reduced laterally, the hind femur may be advanced far forward to form
not more than a 45° angle with the longitudinal axis of the body.

135

b

Fig. 54. Haliplus sp. ; a—ventral view of abdomen; b—metasternum from inside;
st2—second sternite.

The interrelationships of the Triaplidae, Haliplidae and the remaining
Adephaga are presently obscure. It is unclear whether the Triaplidae may be
considered the ancestors of all Adephaga in generai, or whether they, together
with the Haliplidae, form a sister group to the remaining Adephaga. The first
assumption is supported by the very widely occurring large femoral plates
among the early Adephaga. Their occurrence is apparently even more exten-
sive than has been described. Slender femoral piates are seen usually with
great difficulty in fossii remnants, and if not searched for specifically, they
can be easily overlooked in all specimens, except when the abdomen is
severed. Naturally this type of structure will only be included in a description
when its existence is beyond doubt.

Direct evidence in support of the second assumption is lacking. There is
no obvious synapomorphy between Haliplidae and Triaplidae if the large
femoral plates of the hind coxae are not considered synapomorphic. These
structures are unknown in the Archostemata. Such large plates are absent in
other ancient Adephaga (although less developed plates are fairly common
in both the Hydradephaga and Geadephaga), they are generally absent in the
present-day Adephaga. It seems natural that this is a secondary character.
Unfortunately, nothing can be definitely said about the functional utility of
these structures. They can be viewed as a device to prevent the lowering of
the legs during paddling movements in forms having a spherical coxo-
trochanter articulation of the hind legs. Later on, the femoral plates become
reduced with the development of the hinged articulation. However, paddling
movements in all lower Adephaga, including the Haliplidae, are still highly
imperfect; in fact, they do not swim but walk in water. Nevertheless, the
articulation of the trochanter and coxa is already hinged in the Haliplidae.

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136

Of course, we know that the Triassic Triaplidae cannot be considered
the direct ancestors of all Adephaga. Quite typical Mesozoic Adephaga are
observed together with them, primarily not differing even from the Late
Mesozoic forms. They are represented by both terrestrial and aquatic forms,
as far as can be judged from their external appearance. Apparently their
separation into the Geadephaga and Hydradephaga had already taken place.
Earlier a definite assumption on the origin of the Geadephaga from the
Hydradephaga was expressed. However, now when the Haliplidae is clearly
contrasted to both, it appears less tenable. It is not clear if the ancestor of the
Geadephaga was a member of the Hydradephaga or whether both developed
independently from forms closely related to the Triaplidae, while synapo-
morphy of the abdominal base evolved independently in them. The first
theory seems preferable: the apomorphic structure of the abdominal base
probably arose once in forms closely related to Necronectulus, and both the
Geadephaga and dytiscoid Hydradephaga were derived from them.
Gyrinidae appeared almost simultaneously with both. Already in the Late
Triassic, there existed forms similar to Triadogyrus, which seem to be the
direct ancestors of whirligigs. It is impossible to consider them ancestors of
dytiscoids since their femoral plates are completely reduced whereas such
plates are present in the lower Coptoclavidae.

At present, of all the groups of Mesozoic Adephaga we know the most
about dytiscoids. Aerial respiration of larvae through the spiracles of the
eighth abdominal segment is primarily a characteristic feature of this group.
Already in the known ancient forms from the Jurassic there is compact
segmentation of the larval abdomen. The ninth and tenth segments are
reduced and the only functional spiracles of the eighth abdominal segment
are positioned terminally in the middle of the posterior margin of the tergite.
The rather large, bristle-bearing urogomphi were used for holding on to the
surface film of water during respiration. Benthic (Angaragabus) and rather
specialized nektonic larvae (Parahygrobia, Stygeonectes) with natatory hairs
on their legs, are observed among the Jurassic dytiscoid larvae. It is notewor-
thy that Angaragabus already had suctorial mandibles with an internal canal
and lacking a retinaculum, whereas the mandibles of Parahygrobia still had
the primitive general form without a canal. This suggests that they are a
prototype for the modification of mandibles of all Hydradephaga in general.
The most primitive adult dytiscoids had a structure close to that of Necronec-
tulus.

In the Mesozoic, dytiscoids were most widely represented by Cop-
toclavidae. Their remains are found in many of the Jurassic and Cretaceous
sites of Asia, Europe and Africa. Judging from the eyes, which were com-
pletely separated into upper and lower, the adult beetles were pleustonic
predators or scavengers. The legs were still not perfected as paddling ap-
paratus. The most primitive Coptoclavidae (= Necronectinae), had narrow

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tibiae and tarsi with natatory hairs which closely resembled the legs of
Hygrobia or Haliplus. Most Necronectinae still had rather large femoral
plates. Their larvae (Stygeonectes) were active nektonic predators and in
general still poorly differentiated in structure from the most primitive neck-
tonic dytiscoid, Parahygrobia. However, the main characteristics of the
coptoclavid larvae can already be seen in them. Their mandibles are of the
cutting type, the forelegs have begun their modification into raptorial legs,
and the middle and hind legs into natatory appendages.

The more advanced Charonoscaphinae are known exclusively from the
Late Jurassic in which only adult beetles have been found. They clearly
demonstrate considerable advancement in their adaptation toward swim-
ming. Their body shape became more streamlined; middle and hind tibiae
broadened and bore natatory hairs. The femoral plates were fully reduced;
the medial part of the coxa noticeably projected above the body plane from
the place of leg attachment so that the legs could move forward freely. The
integument in the Charonoscaphinae was distinctly weak and through it the
internal structures could be examined in detail in fossil impressions.

The Early Cretaceous Coptoclavinae particularly stand out in their
specialization; they are as yet represented by a single species, Coptoclava
longipoda Ping, which was nevertheless widely distributed in the Early
Cretaceous of East Asia. The adult beetles were large pleustonic forms with
extremely thin integument like that of the large present-day whirligigs in the
genus Dineutes. In impressions, details of the internal structure can often be
studied far more easily and conveniently than the external appearance of the
beetle (Ponomarenko, 1975). The legs of Coptoclava resemble the legs of
whirligigs. The forelegs are raptorial while the middle and hind legs are
natatory with broadened segments. An oar-like surface is formed by the very
broad segments of the tarsi and tibiae which lack natatory hairs. Although
Coptoclava have the most specialized legs among the dytiscoids, they were
apparently surpassed by the effectiveness of the rowing apparatus of whir-
ligigs.

Coptoclava larvae changed less in comparison with Stygeonectes than
did adult Coptoclava in comparison with Necronectes. Here we see essen-
tially the same structural features but somewhat more advanced. An extra
denticle developed on the mesal surface of the mandible, the raptorial
forelegs and natatory middle and hind legs became more specialized, the
urogomphi became longer with far more consolidated segments, and the main
tracheal trunks (used as hydrostatic device) became more powerful. Overall,
in Coptoclava both the larvae and adults are undoubtedly among the oldest
and most unusual Mesozoic insects known to date.

Other dytiscoids are represented in the Mesozoic only by the [pre-
viously] discussed larvae of Angaragabus and Parahygrobia and by the adult
Liadytes, for which a special family was suggested earlier in the text. Liadytes

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and Angaragabus were described as representatives of Dytiscidae in the
broad sense since it was proposed that the adult and larva were possibly of
one and the same species (Ponomarenko, 1963, a, b). New data neither
conclusively prove nor refute this assumption but perhaps lend support to it
since new structural features have been detected in the adult beetles more
closely relating them to the benthic dytiscoids of Noteridae and Phreato-
dytidae to which Angaragabus is also most closely related. The Liadytidae
have changed very little throughout the entire period of their documented
history. Their few representatives, known from the Middle Jurassic and the
end of the Early Cretaceous, could be placed in the same genus. This forms
a sharp contrast with the history of the Coptoclavidae which underwent
considerable morphological evolution during the same period. The
Liadytidae remained slow-moving forms. Their short hind coxae and long
tibiae suggest a low stroke frequency of the hind legs and slow swimming.

The remaining dytiscoid families are almost unknown from the
Mesozoic. The most ancient representatives of the family Dytiscidae have
been found in the Upper Cretaceous deposits of South Kazakhstan. Highly
fragmentary remains have been found here almost undoubtedly belonging to
a beetle combining the salient features of the most primitive tribes of the
subfamilies Colymbetinae and Dytiscinae. None of the other families of
dytiscoids has been found in the Mesozoic. Their absence provides further
evidence against the assumption that the Hydradephaga originated from
forms near Hygrobiidae or Amphizoidae. In fact, many primitive features are
present in the structure of the adult beetles of these families, but the special-
ized structure of the larvae—with reduced terminal abdominal segments—
does not permit the derivation of other Hydradephaga from them, as was done
by Crowson (1955). With regard to Amphizoidae, Bell presents the valid
argument (1966) that these highly advanced dytiscoids adapting to swift
mountain stream lost their ability to swim. The gill breathing larvae of
Hygrobiidae are more modified than the larvae of other dytiscoids. All these
families, together with Noteridae and Phreatodytidae, may have originated
from forms with adults similar to Liadytes, and larvae combining the most
primitive features of Parahygrobia and Angaragabus.

The Mesozoic history of whirligigs differs significantly from that of the
other Adephaga and is more comparable to the evolution of the Polyphaga.
The Mesozoic whirligigs do not form a single group which may be contrasted
with Cenozoic whirligigs. The phylogenetic lines of extant whirligigs are
clearly traceable deep into the Mesozoic, and the main evolutionary trends
remain the same in the Mesozoic and Cenozoic. Of all the extant Hydra-
dephaga, whirligigs form a unique group widely distributed in the Mesozoic;
the whirligigs of the first half of the Jurassic did not differ essentially from
those of the later period. At the same time, Avitortor are the most primitive
of the known whirligigs, fragments of which were found in the Neocomian

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deposits. The Mesozoic evolution of the whirligigs was very patchy. Forms
with a large number of plesiomorphic features persisted for a long time and
distinct evolutionary stages cannot be distinguished in their Mesozoic his-
tory. It is noteworthy that even in the Upper Cretaceous no representatives
of extant families of the whirligigs have been found, although indirect
indications suggest that they must have appeared right at the middle of the
Cretaceous (Ponomarenko, 1973).

Different views exist on the origin and affinities of the whirligigs: from
the contention that whirligigs are the most primitive of all beetles (Bradley,
1930, based on structural similarities of whirligig and sialid larvae) to the
contention that the whirligigs are affiliated with the second wave of adaptive
radiation of the Hydradephaga and were derived from the lower dytiscoids
(Crowson, 1955, 1960). As already stated, the second contention is totally
unrealistic. Of course, even the first contention is unrealistic, although it is
difficult to view the similarity in the gills and anal hooks of whirligig and
sialid larvae as merely the result of convergence. Undoubtedly, here we have
acase, if not of direct inheritance then, at any rate, of the result of activation
of similar inherited potentials. A similar structure of abdominal gills can also
be seen in the diving beetle Coptotomus and the neuropteran Sisyra. In the
development of these structures, the retention of embryonic abdominal legs
in the larvae is obviously a result of heterochrony. Unfortunately, due to our
poor understanding of the evolution of early oligoneopteran larvae this
problem has not been resolved satisfactorily.

The Late Triassic Triadogyrus, of which we unfortunately have only a
single incomplete impression, seems to be a form quite close to whirligigs.
It had a rather long mesosternum; large, obliquely transverse middie coxae;
hind coxae typical of primitive whirligigs, transversely triangular in shape
with curvilinear forward protruding ridge separating the raised and sub-
merged parts of the coxa; the place of articulation of the middle and hind
coxae has trochanters similar to those in impressions of whirligigs. Also, the
abdomen of Triadogyrus has six visible sternites as is usual for the Adephaga
and not seven as in whirzligigs in which the metepisterna extend to the middle
coxal cavities. Unfortunately, the prothorax and head of Triadogyrus have
not been preserved and hence the structure of its eyes and antennae is not
known.

The aforementioned basic facts of the Mesozoic evolution of the
Hydradephaga correspond best with Bell’s view (1966) on the three evolu-
tionary lines of the Hydradephaga—Haliplidae, dytiscoids, and whirligigs—
and not with Crowson’s view (1960) on two adaptive radiations of
Hydradephaga—the earlier (Haliplidae, Amphizoidae and Hygrobiidae) and
the later (Dytiscidae, Noteridae and Gyrinidae)—originating from primitive
dytiscoids. However, contrary to Bell’s view, these phylogenetic lines were
probably not independent paths in the derivation of aquatic forms from

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terrestrial, but independent paths in the perfection of the aquatic mode of life
already acquired by acommon ancestor somewhere close to the boundary of
the Permian and Triassic. The evolutionary paths followed by these three
groups during the Mesozoic and Cenozoic are totally different.

The Haliplidae changed very little, hardly adapting to active swimming.
In fact, the integumentary respiration of their larvae did not permit a size
increase because of the more favourable volume to surface ratio possessed
by smaller forms. The Haliplidae apparently were never an abundant and
diverse group, and despite their relatively high population density in present-
day water bodies, their feeding habits did not change during the transition
from the Mesophytes to the Cenophytes. Their larvae feed on Charophyta
and filamentous green algae which were typical plants of the Mesozoic water
bodies.

Dytiscoids were the most diverse and abundant of the Hydradephaga in
the Mesozoic as well as in the Cenozoic, represented almost exclusively by
extinct families; the geochronology of representatives of the then extant
dytiscoid families begins only at the end of the Mesozoic (in the Late
Cretaceous they were already affiliated with the Cenophytes). They may
have appeared even earlier but were so rare that their remains eluded us.

Whirligigs are the only present-day family widely distributed in the
Mesozoic. The number of whirligig genera known from the Mesozoic is
nearly equal to that in the extant fauna. This is the first instance when an
adephagan family not distinctly relictual at present, had such a wide and
diverse representation in the Mesozoic. The time of origin of the extant
subfamilies of whirligigs was apparently the Middle Cretaceous at the
boundary between the Mesophytes and Cenophytes; however, even in the
Upper Cretaceous there were still representatives of the Mesozoic groups of
whirligigs. The Mesozoic whirligigs do not form a distinct taxon which could
be contrasted with the Cenozoic. On the contrary, individual representatives
of both eras clearly belong to the same lines which had already diverged in
the Mesozoic. The Mesozoic whirligigs are distinguished not so much by
structural primitiveness, as by a combination of features not present in the
Cenozoic forms.

The Geadephaga undoubtedly are a more compact group than the
Hydradephaga. They form a single phyletic line since the processes of
morphological advancement are similar in the individual branches of the
Geadephaga. Even morphologically they can be considered a fairly unified
group despite the presence among them of phytophages, commensals, social
insects and even parasites along with the more widespread predators. If minor
adaptive modifications—at times reaching a very high level of specializa-
tion—are not considered, the basic structural plan remains the same. The
Rhysodidae occupy the most isolated position. Unfortunately, the study of
fossil material has not yet thrown any light on their interrelations with other

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beetles. The earlier view (Ponomarenko, 1968a) of their affinity with the
Mesozoic forms close to the Necromera described by Martinson (1926) is
yet to be confirmed. Therefore the Rhysodidae have been excluded from
consideration in this text.

The Mesozoic Geadephaga are a fairly unified group, and distinct from
the extant Geadephaga. Based on the character of the hind coxae completely
separating the metasternum and abdomen, or extending to the sides only up
to the lateral corners of the metasternum, they are almost since inception
divided into the Trachypacheidae and Carabidae. Forms which cannot be
reliably included in one family or the other are very rare, although in the
Mesozoic trachypacheids the hind coxae are usually distinctly reduced
laterally unlike the extant Trachypachys. A study of the Mesozoic
Geadephaga confirms the isolation of the Trachypacheidae and their place-
ment in a distinct separate family. However, the main evolutionary structural
transformations of beetles in these two families are similar. The parallel
evolution of these families led to a situation, where almost all researchers
placed the present-day Trachypacheidae in Carabidae. In members of both
families the femoral plates were strongly reduced and the lateral walls of the
middle coxal cavities became consolidated so much so that in the extant and
some Mesozoic forms, the metepisterna and then the mesepimera ceased to
reach the middle coxal cavities. In many advanced forms there is a charac-
terisitc modification in the shape of the prosternum from posteriorly broaden-
ing to narrowing before the base. Many Mesozoic ground beetles have a
similar body shape and resemble the Amphizoa or Metrius in this respect.
Apparently their mode of life was far less varied than in the extant carabids.
However, it is rather difficult to be sure about it.

The Mesozoic Trachypacheidae are characterized by the inclusion of the
metepisterna into the lateral walls of the middle coxal cavities and may be
placed in a separate subfamily Eodromeinae. In addition, most of them have
characteristically rather large femoral plates and freely articulating ab-
dominal sternites such that the terminal ones are often telescopic. The
Eodromeinae are known from the Jurassic and Early Cretaceous. The fossils
of their latest representatives found in Aptian-Elbian deposits have large
femoral plates and resemble the Early Jurassic forms.

In the Mesozoic, the Carabidae apparently varied widely both mor-
phologically and ecologically. A large proportion of them belong to strictly
Mesozoic taxa. They all seem to belong to Isochaeta, but in fossil remains it
is difficult to determine the structure of the forecoxae that differentiates
Isochaeta and Simplicia. In the subfamily Protorabinae, the metepisterna
characteristically extend to the middle coxal cavities. These forms are found
in the Jurassic and Neocomian. In external appearance they are very similar
io the Eodromeinae and probably differed little from them ecologically.

142

From the Mesozoic, two markedly different, rather primitive genera of
Protorabinae are known. These are Mesorabus from the Late Jurassic and
Conjunctia from the Early Cretaceous. The total displacement of the pleural
sclerites from the lateral walls of the middle coxal cavities is a characteristic
feature of these genera. The entire coxal cavity in them is formed by the
meso- and metasterna. In this diagnostic character they show parallelism with
the most advanced carabid subfamily Harpalinae. They differ in external
appearance from other Mesozoic ground beetles. Their body is cylindrical,
elongate, with a noticeable constriction between the pronotum and elytra;
externally they resemble ground beetles of the subfamily Scaritinae. Because
of their closed forecoxal cavities and apical spurs of the foretibiae, they can
be included only in Isochaeta or lower Simplicia. Consequently, they con-
stitute an aberrant Mesozoic group forming an early analogue of the sub-
sequently evolved most advanced present-day ground beetles. The more
primitive subfamilies of the extant beetles such as paussinae s.1. and Cara-
binae were not found earlier in the Middle Cretaceous. Yet already by the
Mid-Late Cretaceous, ground beetles externally strongly resembling the
specialized molluskophages of the subfamily Carabinae, and even the first
representative of Harpalinae are found. Thus here, as among the dytiscoids,
we observe a variation in beetles in the Middle Cretaceous; the Mesozoic
forms occur up to the Middle Cretaceous but subsequently the Cenozoic
forms are found.

SUBORDER POLYPHAGA

Among all ancient beetles, those of the suborder Polyphaga have been
studied the least. These ancient beetles are known almost exclusively from
the remains of adult insects, whereas it is known that evolutionary transfor-
mations in the structure of the larvae are especially characteristic of this
suborder. The main groups of this suborder are established on the basis of
larval adaptations. Therefore affiliation of a beetle to many of the major
groups of Polyphaga cannot be determined from the fossil adult. Beetles of
this suborder often lead a cryptic mode of life and their imaginal life span is
generally shorter than that of other beetles; they rarely end up in tapho-
cenoces. The adult beetles develop frequently through regressive metamor-
phosis, they lose many features characteristic of adults and come to resemble
pupae. Beetles of different groups resemble one another. The study of
Polyphaga is rather difficult due to the diversity of this group, comprising
about 150 families and several hundreds of thousands of species. This renders
the classification even of the extant forms unstable and makes the study of
their fossils an extremely difficult undertaking. Apparently, our information
on ancient Polyphaga will remain fragmentary in the near future, and a

143

coherent picture will not emerge. Actually, at present we study beetles which
are marked by specific structures of unique external appearances which set
them apart from all the diverse Mesozoic Polyphaga. Staphylinidae,
Elateridae or* Rhynchophora are such types of beetles. Thus, highly special-
ized forms, which have ceased to evolve any further, are taken up for study
while generalized forms, more interesting from the point of view of
phylogeny, remain unstudied since their taxonomic position cannot be ascer-
tained. It would be worthwhile to take a select number of these more
generalized forms which are of particular interest in one respect or another,
and formally describe them as a group having an unclear systematic position.

POLY PHAGA INCERTAE SEDIS

Genus Peltosyne Ponomarenko, gen. nov.

Genus name coined from ‘pelta’ (Greek)— shield, and genus Ademosyne.
Type species. P. triassica, sp. nov. Triassic of Soviet Central Asia.
Description. Rather large flat beetles. Head large, eyes lateral, gular

plate wide. Pronotum transverse, not longer than head, anteriorly emarginate,

its lateral margin sharp, pleural suture distinct. Prosternum with rather wide
process slightly broadening beyond coxae, its length more than that of
prothorax anterior to coxae. Forecoxal cavities posteriorly closed or almost
closed by lateral projections. Forecoxae transverse, rectangular, with dis-
tinct trochanters. Mesothorax rather short, mesosternum shorter than
mesopleuron, in the middle with a depression for apex of prosternal process,
and with traces of transverse suture. Mesepisterna much smaller than mese-
pimera, remote from middle coxal cavities. Middle coxae small, set apart,
with distinct trochanters. Metathorax large, metasternum slightly narrowing
anteriorly, transverse metasternal suture distinct laterally, medially slightly
curved backward and fusing with anterior margin of hind coxae. Metepister-
na almost not broadening anteriorly, not reaching middle coxae. Metepimera
somewhat broadening posteriorly, but not forming distinct mesal tonge. Hind
coxae short, contiguous, posteriorly excavated but without distinct femoral
plates. Abdomen with five visible sternites. Elytra with punctate grooves.

Integument with dense and sharp punctation, on the elytra rugose.

Species composition. Monotypic genus.

Taxonomic position. The species described here is the most ancient of
the well preserved Polyphaga known to date. It is still very similar to
Ademosynidae, the representative of Archostemata very close to Polyphaga,
particularly in its morphology. At that time, Peltosyne almost certainly
already possessed a cryptopleural prothorax, and metepisterna not reaching

*So given in the Russian original. Since Elateridae are not a part of Rhynchophora “or” should
be substituted with “and ”—General Editor.

144

the middle coxal cavities. It is difficult to define the position of Peltosyne in
Polyphaga. The structure of the base of abdomen and wings is not known.
Almost any attempt to combine it with any series of superfamilies (Crowson,
1955, 1971) in Polyphaga will amount to a symplesiomorphic unification.
Peltosyne is on the whole a rather generalized type of polyphagan beetle and
may be considered very close to the ancestral type of this suborder. Of course,
it cannot be considered the direct ancestor of all Polyphaga. Even more rare
representatives of other groups of Polyphaga, including such advanced forms
as Rhynchophora, are found in the same deposits. It would not be advisable
to exclude Peltosyne from any one of the large taxa of extant beetles or to
consider it representative of a special high ranking taxon. It is more correct
to describe it at present as Polyphaga incertae sedis and use the level of
development of Polyphaga during the Triassic for its characterization.

It is premature to establish to taxon of the rank of family on the basis of
the genus Peltosyne at this stage (= Peltosynidae fam. nov.), in view of the
inadequacy of the data on this form which is known from fragmentary
remains.

Peltosyne triassica Ponomarenko, sp. nov.
(Plate IX, Photo 1; Figure 55)

Species name coined from Triassic.

Holotype. No. 2240/278, PIN, impression of beetle without antennae
and legs. Southern Ferghana, Kirgiz SSR, Osh oblast, Batken region,
Madygen area (Dzhailyaucho site). Triassic.

Material. Holotype and a fragment of elytron, specimen No. 2240/230
from the same site.

Description. Head tapering from base anteriorly, genae very short,
temples about half the length of eyes. Head without distinct sutures dorsally.
Mandibles rather large, fully half the length of head capsule. Gular plate
broadening in anterior half. Prosternum anterior to coxae a little shorter than
them. Mesosternum not longer than middle coxae, mesepisterna half the
length of mesepimera. Metasternum two-and-a-half times longer than middle
coxae, width of its anterior margin two-thirds the posterior. Hind coxae
shorter than middle coxae. Basal four abdominal sternites equal in length,
last sternite twice as long, basal width of last sternite half of abdominal base,
abdomen narrowing from base of third sternite.

Dimensions. Body length 11 mm, width 6 mm, elytral length 8 mm.

Remarks. The fossils from Dzhailyaucho site underwent strong post-
fossilization distortion (Ponomarenko, 1969). Hence the size of sclerites can
be compared only along one of the body axes of the beetle.

145

eo)
\A
be
pees (

a b

Fig. 55. Peltosyne triassica, sp. nov.; holotype PIN No. 2240/278: a—dorsal
view; b—ventral view. Dzhailyaucho, Tnassic.

INFRAORDER STAPHYLINIFORMIA LATREILLE, 1802

Family HYDRAENIDAE Mulsant, 1844
Formal genus “Ochthebiites*”

“Ochthebiites” altus Ponomarenko, sp. nov.
(Plate IX, Photo 2; Figure 56)

Species name coined from ‘altus’ (Latin)—ancient.

Holotype. No. 3000/917, PIN, impression of a beetle without a large part
of legs. Trans-Baikal, Buryat ASSR, Mukhorshibir region. Novospasskoe
village (Novospasskoe site). Middle Jurassic, Ichetui series.

107 Material. Holotype and specimen No. 3000/925 from the same site.

Description. Very small beetles. Body rather wide, flat, robust. Length
of head slightly shorter than average width, widest near eyes, from them
narrowing anteriorly and toward base. Antennae noticeably longer than
palps, their five apical segments moniliform, basal segments short. Labrum

*Here and elsewhere in this section the genus name is incorrectly given as Ochtebiites in the
Russian original—General Editor.

146

Fig. 56. “Ochthebiites” altus, sp. nov.; a—holotype PIN No. 3000/917;
b—paratype PIN No. 3000/925. Novospasskoe, Jurassic.

with rather deep emargination at middle. Segments of palps short, last
segment broadly lanceolate, twice as long as wide. Width at anterior margin
of pronotum 1.7 times its length, narrowing beyond anterior third, posterior
margin approximately two-thirds the anterior; anterior corners rounded.
Prosternum noticeably shorter than forecoxae. Mesosternum almost equal in
length to rounded middle coxae, latter converging. Length of metasternum
two-fifths its width at posterior margin, width at anterior margin two-thirds
the posterior margin, latter angularly projecting backward between coxae.
Hind coxae extending backward along mid-line, overlapping almost entire
first visible abdominal sternite. Abdomen with seven visible sternites, two
terminal ones sharply tapering, last sternite noticeably longer than wide.
Abdominal segments apparently telescoping. Legs rather short, femora
weakly uniformly thickened, hind femora a little longer than middle femora.
Tibiae slender, linear. Foretarsi half and hind tarsi two-thirds the length of
corresponding tibiae, four* basal segments of tarsi approximately equal in
length to the fourth*. No ornamentation visible on head and pronotum. Elytra
with blunt grooves.

Dimensions. Body length 2.2—2.4 mm, width 1.2 mm; elytral length 1.6
mm.

Taxonomic position. Based on the structure of the distal part of anten-
nae, palps, legs and abdomen, these fossils almost certainly belong to the
family Hydraenidae whose members differ very slightly from the extant

*Translation correct; number of segments needs verification—General Editor.

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147

genus Ochthebius and closely related forms. Proximity to Ochthebius is seen
in the short palps, and convergent middle coxae. However, these very small
beetles are not sufficiently well preserved for comparison with the extant
genera in which the structures of antennal base and genitalia are charac-
teristic. These structural details are not known in the extinct forms, and since
itis impossible to formally prove that they do not belong to a particular extant
genus, they are placed in a formal genus, “Ochthebiites”. The sole specific
feature of the described forms is apparently the highly transverse pronotum,
but this distinguishing character is hardly sufficient for establishing an
independent genus. It is reaily remarkable that in the Middle Jurassic, forms
existed which differed very slightly from the extant hydraenids.

“Ochthebiites” incertus Ponomarenko, sp. nov.
(Figure 57)

Species name coined from ‘incertus’ (Latin)—uncertain.

Holotype. No. 2997/1898, PIN, impression of almost entire beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka villag2 (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype. ,

Description. Very small beetle. Body rather wide, robust, flat. Head
shorter than its width, tapering anteriorly. Length of prothorax two-thirds its
width, pronotum widest in anterior third, tapering toward base and apex.

oN

Fig. 57. “Ochthebiites” incertus, sp. nov.; holotype PIN No. 2997/1898, Karatau,
Upper Jurassic.

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148

Abdomen with seven visible sternites, last two tarsal segments slightly longer
than remaining [basal] four together. Elytra with punctate grooves.
Dimensions. Body length 2.9 mm, 1.4 mm; elytral length 2.0 mm.
Taxonomic position. The beetle is extremely poorly preserved, hence its
taxonomic position could not be formally confirmed. However, the structure
of abdomen and tarsi suggests its inclusion in the family Hydraenidae. It is
very similar to the species described above, differing only by somewhat
larger dimensions, shape of prothorax and more distinct punctures on elytra.

Family HYDROPHILIDAE Leach, 1815*

2K
Genus Mesospercheus Ponomarenko, gen. nov.

Genus name coined from Mesozoic, and genus Spercheus.

Type species. M. tarsalis, sp. nov. Lower Cretaceous of Trans-Baikal.

Diagnosis. Small flat beetles. Head with Y-shaped suture, clypeus trun-
cate, labrum concealed under it, palpal segments short. Pronotum transverse,
laterally rounded, [underside of] prothorax with grooves for antennae, and
with mesal projection beyond forecoxae. Abdomen with five sternites. Last
tarsal segment distinctly longer than four preceding together. Large em-
podium absent.

Species composition. Besides the type species, the genus includes poor-
ly preserved remains of similar hydrophilids from the Jurassic Ichetui series
of western Trans-Baikal.

Taxonomic position. Among the hydrophilids, Mesospercheus occupies
a fairly isolated position. In the long last segments of the tarsi, it shows
affinity with Hydrochus and Spercheus which are often regarded as repre-
sentatives of separate monotypic families. It differs from Hydrochus in the
absence of a freely articulating labrum and absence of transverse ridges on
abdominal sterniies; it more closely resembles Spercheus in these characters.
However, specific apomorphic characters of Spercheus, such as the densely
bristled empodium, are definitely absent in Mesospercheus, and the clypeus
is devoid of an emargination. It has not been possible to study the structure
of antennae in the available fossil material. Thus, proximity to Spercheus is
also based on symplesiomorphy and their taxonomic unification is hardly
advisable. Apparently, Mesospercheus is a primitive form, closely affiliated
to the common ancestor of Spercheus, Hydrochus, and Hydraenidae.

*Family Hydrophilidae is sensu lato considered to include Hydrochus and Spercheus.

**Given as Mesosperchus in the Russian original. The spelling of this genus needs to be
confinmed. If it is a combination of mesozoic and Spercheus the correct spelling ought to be
Mesospercheus. Compare also p. 109 of Russian original—General Editor.

110

149

Mesospercheus tarsalis Ponomarenko, sp. nov
(Plate IX, Photos 3, 4; Figure 58)

Species name coined from ‘tarsus’ (Latin)—sole of foot.

Holotype. No. 3015/367, PIN, impression of a beetle lacking antennae
and large part of legs. Trans-Baikal, Chita oblast, Balei region, right bank of
Unda river 2 km upstream from Zhidka settlement (Unda site). Lower
Cretaceous, Aptian-Albian ? Balei series.

Material. Holotype and impression of an almost entire beetle, specimen
No. 3063/116 from the same site.

Description. Head almost triangular, narrowing from base, its length
slightly less than width. Stem of Y-shaped suture not exceeding fork in
length. Clypeus large, its anterior corners rounded, middle part truncate.
Gular plate narrow. Palps rather short, their apical segment longest, twice as
long as wide. Pronotum short, rounded laterally, anteriorly emarginate, its
length two-fifths the width. Prosternum short, much shorter than projecting
forecoxae. Mesosternum shorter than middle coxae, mesepisterna slightly
shorter than mesepimera, suture between them almost transverse. Middle
coxae rounded, converging. Metasternal length two-fifths its width at
posterior margin, width of anterior margin two-thirds the posterior.
Transverse metasternal suture visible only laterally, in middle third fused
with anterior margins of hind coxae. Metepisterna slightly broadening
anteriorly, their width at anterior margin one-third the length. Hind coxae
oblique, extending backward and broadening along midline of body. Ab-
domen tapering from base of second visible sternite, basal width of last
sternite half of abdominal base, all five sternites subequal in length. Legs
short, femora scarcely extending beyond lateral body margins. Forefemora
rather strongly thickened, remaining femora slightly thickened. Tibiae
uniformly broadening from base to apex and curved. Foretibiae three-fourths
the length of middle tibiae, half the length of hind tibiae. Tibiae with spines
laterally, foretibiae with strongly curved apical spur. Last tarsal segment
one-and-a-half times longer than the equal remaining segments. Elytra with
distinctly deep grooves, scutellar groove long, joining sutural groove.

Dimensions. Body length 4.0-4.1 mm, width 1.7—1.9 mm; elytral length
2.8 mm.

Comparison. Differs from the second species in shorter pronotum which
is more strongly and angularly emarginate at anterior margin.

Mesospercheus notatus Ponomarenko, sp. nov.
(Plate [X, Photo 5; Figure 59)

Species name coined from ‘notum’ (Greek)—dorsal shield.

150

Holotype. No. 3000/923, PIN, impression of beetle lacking head and a
large part of legs. Trans-Baikal, Buryat ASSR, Mukhorshibir region,
Novospasskoe settlement (Novospasskoe site). Middle Jurassic, Ichetui

series.

c t-s

b ey

Fig. 58. Mesospercheus tarsalis, sp. nov.; a—holotype PIN No. 3015/367; b,

c—paratype PIN No. 3063/116: b—dorsal view, c—ventral view. Unda, Lower
Cretaceous.

111

151

Material. Holotype and impression of abdomen and elytra of beetle,
specimen No. 3000/912 from the same site.

Description. Width of pronotum 1.8 times the length, anteriorly with
shallow rounded emargination, rounded laterally, maximum width in
posterior third; anterior margin markedly narrower than posterior. Underside
of prothorax with distinct groove for antennae near anterior angles. Proster-
num small, shorter than projecting forecoxae. Middle coxae convergent,
slightly transverse, oblique. Metasternum slightly narrowing anteriorly, its
length half the width at posterior margin, latter angularly projecting medially,
extending between coxae. Hind coxae almost transverse, only medially
extending backward. Abdominal sternites approximately equal in length.
Legs short, femora scarcely extending beyond lateral body margins. Hind
tibiae slightly longer than middle tibiae. Middle and hind tibiae with spines
laterally. Hind tarsi half the length of tibiae, last segment one-and-a-half
times longer than basal segments together. Elytra with distinct grooves,
extending to outer margin apically.

Dimensions. Body length about 6.5 mm, width 2.0 mm; elytral length
5.2-5.5 mm, length of pronotum 0.8 mm.

Comparison. Differs from type species in sormewhat larger dimensions,
and longer, weakly anteriorly emarginate pronotum.

Remarks. Both available fossils of the beetle are fragmentary. Hence it
has been affiliated to the genus Mesospercheus and to hydrophilids in general

Fig. 59. Mesospercheus notatus, sp. nov.; a—hoiotype PIN No. 3000/923; b—
paratype PIN No. 3000/912. Novospasskoe, Jurassic.

112

52

only on the basis of similarity with the preserved part of the body of M.
tarsalis.

Genus Mesydra Ponomarenko, gen. nov.

Genus name coined from ‘mesos’ (Greek)—middle, and ‘hydor’ (Greek) —
water.

Type species. M. elongata, sp. nov. Lower Cretaceous of Trans-Baikal.

Diagnosis. Small, elongate, cylindrical beetle. Head transverse, with
distinct Y-shaped suture. Eyes with large ommatidia. Antennae short, with
compact three-segmented oval club and monoliform short segments behind
it. Pronotum transverse, with rounded sides. Elytra at shoulders scarcely
wider than pronotum. Scutellum large, triangular. Wings with well-develo-
ped venation. M + Cu cell present, bending at midwing. Abdomen long, with
six visible sternites, last sternite longest. Legs short, four basal segments of
tarsi approximately equal to last segment or shorter. Elytra without distinct
punctate grooves. Sides of pronotum and last abdominal sternite with long
hairs.

Species composition. Monotypic genus.

Taxonomic position. Judging from the antennal structure and complete
wing venation, this species must be included in Hydrophilidae in the broad
sense considered in this work. In the structure of palps and legs it resembles
ordinary Mesozoic hydrophilids. Also, Mesydra has several distinguishing
characters bringing it closer to Hydraenidae: abdomen with six visible
sternites, long hairs on pronotum and abdomen, apical bend of elytra strongly
shifted proximally. However, considering the possible paedomorphic nature
of all these changes, it may be considered a member of Hydrophilidae
whereas the Hydraenidae features developed in it independently.

Mesydra elongata Ponomarenko, sp. nov.
(Plate IX, Photo 6; Figure 60)

Species name coined from ‘elongata’ (Latin)—elongated.

Holotype. No. 1989/2983, PIN, reverse impression of almost entire
beetle with hind wings spread. Trans-Baikal, Buryat ASSR, Eravnin region,
left bank of Vitim river downstream from the mouth of Baisa river (Baisa
site). Lower Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Width of head 1.7 times its length, narrowing from base
anteriorly, temples and genae very short. Y-shaped suture distinct, its stem
shorter than fork, angle between fork almost 90°; fronto-clypeal suture
distinct. Eyes with very large sparse ommatidia, about ten across the width
of eye. Antennal club compact, its length one-and-a-half times the width.

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153

Fig. 60. Mesydra elongata, sp. nov.; holotype No. 1989/2983: a—ventral view;
b—dorsal view of head, and pronotum. Baisa, Lower Cretaceous.

Palps not longer than head, last segment one-and-a-half times longer than
penultimate. Width of pronotum 1.8 times the length, anteriorly weakly
roundly emarginate, posteriorly projecting, anteriorly not narrowing, angles
rounded. Scutellum as a large equilateral triangle, its sides only one-third the
basal width of pronotum. Width of metasternum at posterior margin 2.5 times
its length, almost not narrowing anteriorly, distance between middle and hind
coxae shorter than middle coxae. Thorax and abdomen in different planes,
with a noticeable drop in elevation between. Abdomen tapering from base
of third visible sternite, basal width of fifth sternite two-thirds of abdominal
base, width of the sixth sternite half the abdominal base. Length of last
sternite half its basal width, almost twice as long as remaining sternites.
Forefemora slightly but uniformly thickened, nearly equal in length, tibiae
slightly shorter than them, slightly broadening from base to apex. Foretibia
with large curved spur. Foretarsi slightly shorter than tibiae, last segment
almost one-and-a-half times longer than preceding segments together. Mid-
dle tarsi two-thirds the tibiae, last segment equal to penultimate. Wings
approximately twice as long as elytra. Body with fine indistinct punctation.

Dimensions. Body length 5.6 mm, width 2.3 mm; elytral length 3.3 mm,
wing span 5.8 mm.

154
Genus Mesohelophorus Ponomarenko, gen. nov.

Genus name coined from Mesozoic, and genus Helophorus.

Type species. M. sibiricus, sp. nov. Lower Cretaceous of Trans-Baikal.

Diagnosis. Small, flat beetle. Head large, with distinct Y-shaped suture
on top. Pronotum strongly transverse, trapezoidal, weakiy narrowing
anteriorly, laterally flat, disk with longitudinal grooves. Middle coxae con-
vergent, with keel-shaped intercoxal process. Abdomen with five visible
sternites. Legs short, last segments of fore- and middle tarsi equal to all the
preceding segments [together], last segments of hind tarsi shorter than them.
Elytra with sharp punctate grooves.

Species composition. Monotypic genus.

Taxonomic position. This genus exhibits a distinct similarity with
Helophorus in the characteristics of tarsi, transverse not anteriorly narrowing
pronotum and the structure of middle coxae. In the laterally flat-sided
pronotum with distinct corners, this genus generally differs from all
hydrophilids including Helophorus, it also differs in the structure of elytra.
Mesohelophorus has only three longitudinal grooves on the disk of the
pronotum, and not five as in Helophorus.

Mesohelophorus sibiricus Ponomarenko, sp. nov.
(Plate IX, Photo 7; Figure 61)

Species named after Siberia.

Holotype. No. 3063/841, impression of beetle lacking antennae and
large part of legs. Trans-Baikal, Buryat ASSR, Eravnin region, left bank of
Vitim river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype, incomplete beetle no. 3064/931 isolated elytron
No. 1989/2998 from the same site.

Description. Length of head a little less than width, widest near eyes,
tapering toward base, triangular in front of eyes. Eyes convex. Stem of
Y-shaped suture slightly shorter than fork, angle between fork obtuse. Gular
plate narrow. Antennae difficult to see, apparently of structure normal for
hydrophilids, with two long basal segments followed by three short,
moniliform segments; club oval, three-segmented, wide capsular, thick and
compact. Antennae slightly extending beyond prothorax. Length of
pronotum almost one-third the width at posterior margin, anterior margin
scarcely narrower than posterior, pronotal angles almost right-angled.
Anterior margin weakly roundly emarginate, posterior slightly projecting.
Middle part of pronotum approximately half its width, [taking the form of a]
raised flat dome above the flattened lateral fourths. Longitudinal groove
passing medially through the dome, deepening into a pit at posterior margin

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155

of pronotum. Shallower grooves running on both sides of the dome. Sides of
pronotum margined with narrow lines. Mesosternum scarcely shorter than
middle coxae, mesepimera oblique, their length less than width. Middle
coxae rounded, triangular, closely convergent; long, narrow keel-shaped
mesosternal projection and short broad metasternal process present between
them.

Length of metasternum two-fifths the width at posterior margin, width
at anterior margin two-thirds that at posterior. Metepisterna strongly
broadening anteriorly, hind coxae transverse, length of its raised part reduced
laterally, elongate posteriorly along the mid-line; lateral margin of elongate
medial part emarginate. Abdomen short, not longer than meso- and
metathorax together, narrowing posteriorly from base of last sternite, width
of the latter half of abdominal base. All sternites almost equal in length, freely
articulate. Apices of slightly thickened femora scarcely extending beyond
lateral body margins. Fore- and middle tibiae noticeably shorter than femora,
hind tibiae equal to them. Foretarsi approximately half the length of tibiae,
middle tarsi equal to tibiae, hind tarsi slightly longer. Last segment of hind
tarsus equal to the two preceding. Elytral grooves running parallel to sutural
margin. Scutellar groove long, not shorter than one-third of elytron. Intervals
between punctures in furrows only a little larger than the punctures them-
selves. Integument strongly punctate. Sinciput, occiput and pronotum with
large, coarse punctation, intervals between punctures not larger than punc-
tures themselves, elytra and large part of body ventrally shagreened, under-
side of pro- and mesothorax with rather fine dense punctation.

Fig. 61. Mesohelophorus sibiricus, sp. nov.; holotype PIN No. 3064/841: a—dorsal
view; b—ventral view. Baisa, Lower Cretaceous.

115

156

Dimensions. Body length 3.5 mm, width 1.8 mm, elytral length 2.1 mm.

Genus Paraspercheus Ponomarenko, gen. nov.

Genus name coined from genus Spercheus.

Type species. P. asiaticus, sp. nov. Upper Jurassic of South Kazakhstan.

Diagnosis. Rather large, flat, broad-beetles. Head large, elongate, par-
tially covered by pronotum. Y-shaped suture indistinct. Antennae rather long,
with long, slender basal segments and compact, oval, three-segmented club.
Pronotum transverse, rather strongly narrowing anteriorly, anteriorly emar-
ginate. Forecoxae large, projecting. Mesosternum slightly shorter than mid-
dle coxae. Middle coxae large, slightly separated, almost equal in length and
width, with prominent anterodorsal part. Metasternum strongly transverse.
Hind coxae contiguous. Abdomen with five sternites. Foretibiae with large
curved spur. Last segment of tarsi longer than preceding ones together. Large
empodium absent. Elytra without grooves. Integument with coarse puncta-
tion.

Species composition. One species from Late Jurassic of South Kazakh-
stan and one species from Early Cretaceous of Trans-Baikal.

Taxonomic position. Genus Paraspercheus occupies the most isolated
position among Hydrophilidae s. 1., i.e. hydrophilids having a normal ab-
domen with five visible sternites. Trailing legs with a very long last segment
are observed in only a few small, distinctly relictual forms. In general,
Paraspercheus is related to the largest hydrophilid beetles. It differs marked-
ly from other hydrophilids in possessing very coarse punctation of the
integument, totally lacking traces of grooves on the elytra and also in the
absence of a large empodium; the latter structure is highly characteristic of
many hydrophilids, at least of forms with an enlarged middle tarsal segment.
The empodium is definitely absent in Paraspercheus since if it could be seen
in fossils of small hydrophilids then it ought to be seen in the well preserved
tarsi of Paraspercheus vitimensis. Apparently Paraspercheus is a repre-
sentative of the special Mesozoic line of hydrophilids.

Paraspercheus asiaticus Ponomarenko, sp. nov.
(Plate IX, Photo 8; Figure 62)

Holotype. No. 2997/552 [sic], PIN, impression of almost entire beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of the
Kashkarata river valley, Aulie area in Mikhailovka village (Karatau-Mik-
hailovka site). Upper Jurassic, Karabastau series.

Material. Holotype and impression of one almost entire beetle,
specimen No. 2239/1224 from the same site.

Description. Head slightly longer than wide, slightly narrowing anterio-
rly, with constriction in front of base, length of temples half the eyes, genae

116

157

almost equal to eyes in length. Y-shaped suture weak, its stem longer than
fork, angle between fork obtuse. Gular plate narrow. Clypeus and labrum
strongly projecting anteriorly, labrum long, distinct, degree of its sclerotiza-
tion distinctly second to that of clypeus. Mandibles rather long, strongly
projecting anteriorly. Three basal antennal segments equal in length, first
noticeably thicker than second and third, following segments shorter than
third in length, slightly moniliform; three apical segments forming rather
compact oval, asymmetric club. Length of pronotum half its width, laterally
rounded, narrowing anteriorly and posteriorly, widest at middle. Prosternum
shorter than large projecting forecoxae. Distance between middle coxae
approximately one-third their width. Mesepimera transverse, short. Width of
metasternum at posterior margin 3.5 times its length, width at anterior margin
two-thirds of posterior. Distance between middle and hind coxae less than
the length of middle coxae. Metepisterna slightly broadening anteriorly. Hind
coxae transverse, their raised medial parts extending backward and laterally.
Abdomen longer than meso- and metathorax together, tapering from base of
third sternite; basal four sternites equal in length; last segment one-third
longer, triangular, its basal width half of abdominal base. Legs noticeably
extending beyond lateral body margins. Forefemora almost equal to middle
femora and two-thirds the hind femora. Foretibia slightly shorter than femora,

ma

a b

Fig. 62. Paraspercheus asiaticus, sp. nov.; a—holotype PIN No. 2997/522,
b—paratype PIN No. 2239/1224. Karatau, Upper Jurassic.

158

flat, with grooves and rows of spines laterally. Apical spur of foretibiae
reaching third tarsal segment. Middle tibiae noticeably shorter than cor-
responding femora, narrower than foretibiae. Hind tibiae slender, linear, 1.4
times longer than foretibiae. Foretarsi only slightly shorter than foretibiae,
first segment much shorter than second, last noticeably longer than the
remaining segments together. Middle tarsi not shorter than tibiae, their last
segment scarcely longer than remaining segments together. Hind tarsi ap-
proximately half the hind tibiae, last segment slightly longer than remaining
segments taken together. Claws large, weakly curved. Sinciput, occiput,
pronotum and prosternum with coarse punctation, punctures on meso- and
metasterna smaller and dense, elytra with large scattered punctures.

Dimensions. Body length 14-15 mm, width 8 mm; elytral length 9 mm.

Comparison. Distinguished by pronotum narrowing toward base, longer
head, narrow foretibiae, and absence of distinct ornamentation.

Paraspercheus vitimensis Ponomarenko, sp. nov.
(Plate IX, Photo 9; Figure 63)

Holotype. No. 3064/1071, PIN, incomplete reverse impression of beetle,
only structure of dorsal side and part of legs visible. Trans-Baikal, Buryat
ASSR, Eravnin region, left bank of Vitim river downstream from the mouth
of Baisa river (Baisa site). Lower Cretaceous, Valanginian-Goteriv, Zazin
series.

117 Fig. 63. Paraspercheus vitimensis, sp. nov.; holotype PIN No. 3064/1071. Baisa,
Lower Cretaceous.

O17

159

Material. Holotype.

Description. Oval beetle. Head rather strongly retracted under prono-
tum, noticeably longer than wide, weakly tapering anteriorly. Y-shaped
suture weak, its stem much longer than fork, angle between fork obtuse,
mandibles strongly projecting anteriorly. First to third segments of antennae
long; 4th shorter than them, transverse. Width of pronotum at posterior
margin 1.8 time its length, roundly tapering anteriorly, width at anterior
margin two-thirds the posterior. Foretibiae scarcely shorter than middle
tibiae, two-thirds the hind tibiae, broad and flattened, with longitudinal
groove and two rows of spines; spines of outer row larger and sparser than
inner row. Tibiae with denticles laterally. Spur of foretibiae strongly curved,
equal in length to three basal segments of foretarsi. Middle tibia broadening
from base to apex, with longitudinal groove, laterally serrate, spur of middle
tibiae shorter than two basal tarsal segments. Foretarsi equal to tibiae in
length, first segment shorter than second, last noticeably longer than all
preceding ones together. Middle tarsi equal to middle tibiae, their three basal
segments equal, slightly shorter than fourth segment, together with it equal
to fifth. Claws long, almost straight. Dorsal body surface with coarse punc-
tures, somewhat finer punctures on head. Intervals between punctures
smaller than punctures themselves. Pronotum with dark, transversely oval
spot in front of posterior margin, elytra with dark colored suture and four
longitudinal dark colored bands on the disk laterally.

Dimensions. Body length about 13 mm, width 7.5 mm; elytral length 8.5
mm.

Comparison. Differs from the type species in the more retracted head,
pronotum narrowing from base, and broad forefemora. Probably merits status
of an independent genus.

Family SILPHIDAE Latreille, 1807
Genus Mesagyrtes Ponomarenko, gen. nov.

Genus name coined from Mesozoic and genus Agyrtes.

Type species. Mesagyrtes communis, sp. nov.; Jurassic of Trans-Baikal.

Description. Small elongate-oval beetle with short slender legs. Head
equal in length and width, widest at occiput, whence narrowing anteriorly.
Eyes slightly protuberant, ocelli present on hind corners of sinciput. Anten-
nae extending beyond pronotal base, their distal segments broadening, form-
ing loose club. Segments of maxillary palps cylindrical, last segment only
slightly larger than penultimate, not dilated. Pronotum transverse, narrowing
from middle anteriorly far more strongly than posteriorly, scarcely narrower
than elytra at shoulders, flat lateral margin absent. Scutellum rather large,
triangular. Prosternum short, distinctly emarginate anteriorly, taenioid, much
shorter than forecoxae. Forecoxae projecting, but rather short, with long

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160

narrow lateral process. Foretrochanters virgate. Mesosternum shorter than
middle coxae, mesepimera equal to middle coxae in length. Metasternum
transverse, with distinct transverse and longitudinal metasternal sutures.
Middle coxae slightly transverse, oblique, projecting. Metepisterna not
broadening anteriorly. Hind coxae contiguous, transverse, short. Abdomen
with five visible sternites, last sternite not longer than preceding ones,
apically truncate. Elytra with very blunt longitudinal grooves beset with faint
punctures. Femoral apices scarcely extending beyond lateral body margins,
femora slightly and uniformly thickened, tibiae slender, slightly broadening
toward apex, tarsi with slender cylindrical segments.

Species composition. Monotypic genus.

Taxonomic position. Based on elytra with punctate grooves, the genus
stands closest to tribes Agyrtini and Pterolomini, however, the grooves are
much weaker than in the representatives of these tribes. The presence of ocelli
most closely relates it to Pterolomini though the abdomen with five sternites
puts it closest to Agyrtini. The shape of its body places it intermediate
between these tribes. With only one specimen of this rather isolated form, it
is difficult to define its exact place in the family.

Mesagyrtes communis Ponomarenko, sp. nov.
(Plate X, Photos 1-3; Figure 64)

Species name coined from ‘communis’ (Latin)—common.

Holotype. No. 3000/926, PIN, impression of almost entire beetle. Trans-
Baikal, Buryat ASSR, Mukhorshibir region, north of Novospasskoe settle-
ment (Novospasskoe site). Middle Jurassic, Ichetui series.

Material. Besides holotype, 6 impressions of almost entire beetles,
specimen Nos. 3000/909, 910, 913, 916, 920, 929, impression of head and
pronotum, specimen No. 3000/915 from the same site, reverse impression of
entire beetle, No. 2744/12 from Novaya Bryan’ site, Buryat ASSR, Mukhor-
shibir region, Bryanka river, Ichetui series.

Description. Length of head capsule two-thirds its width, temples
shorter than eyes, genae short. Gular plate longer than wide. Labrum slightly
emarginate apically. Last segment of maxillary palps 1.5 times longer and
slightly thicker than penultimate. First antennal segment noticeably thicker
and longer than second; second half the length of third; third thinnest of all,
almost not broadening towards apex, slightly longer than fourth; fourth and
fifth noticeably broadening toward apex; fifth two-thirds the third; fifth to
tenth segments equal in length, strongly broadening toward apices; apex of
ninth twice as wide as apex of fourth; last segment ovoid, equal in length to
third. Length of pronotum approximately half its width, roundly emarginate
at anterior margin, rounded laterally, width of its posterior margin 1.7 times

119 the width at middle. Length of prosternum one-fifth its width. Lateral process

118

161

of forecoxae not narrower than coxae themselves. Mesosternum half the
length of middle coxae. Width of mesepisterna approximately half of
mesepimera. Middle coxae with sharply truncate and flat lateral part. Length
of metasternum half its width; when middle coxae extending backward, then

Fig. 64. Mesagyrtes communis, sp. nov.: a, b—holotype PIN No. 3000/926: a—dor-

sal view, b—ventral view. Paratypes: c—PIN No. 3000/915, d—PIN No. 3000/913,

e—PIN No. 3000/909, all from Novospasskoe, Jurassic; f—paratype PIN No. 2744/
12, head; Novaya Bryan’, Jurassic.

162

distance between middle and hind coxae less than length of middle coxae.
Length of hind coxae half their width, rather sharply reduced beyond mesal
third. Abdomen tapering from second visible sternite, basal width of last
sternite half of abdominal base, first visible sternite one-and-a-half times
longer than second, remaining sternites almost equal. Length of forefemora
two-thirds the middle femora, half of hind femora, all tibiae slightly shorter
than femora, apices of foretibiae not extending. Tarsi with slender, virgate
segments, slightly less than tibiae in length.

Dimensions. Body length 4.6-5.2 mm, width 1.8-2.3 mm; elytral
length 2.7-3.0 mm.

INFRAORDER SCARABAEIFORMIA
Family SCARABAEIDAE Latreille, 1802

To date, available information on Mesozoic Scarabaeidae was primarily
confined to the single Early Cretaceous Proteroscarabeus Grabau. The
actual affiliation of the remaining forms mentioned here remained absolutely
debatable. It must be noted that one of the presumed scarabaeids-jurassicus
Oppenheim, 1888, was generally described from the impression of a cock-
roach and not a beetle (Ponomarenko, 1971). In the Lower Cretaceous
deposits of Trans-Baikal numerous remains were found whose sublineal
affiliation within the family Scarabaeidae is certain. Almost all the remains
are well preserved intact specimens, and even the structure of hind wings and
position of abdominal spiracles can be successfully studied in some of them.
However, it is a difficult task to establish the precise taxonomic position of
the described forms. The formal affiliation of scarabaeids to a subfamily
requires knowledge of the structure of antennae and position of abdominal
spiracles. These distinguishing characters usually cannot be sufficiently well
studied in fossil remains. The use of indirect, often habitual characters for
such ancient forms is risky. Their comparison with Cenozoic forms therefore
becomes impossible and even the described genera should be considered
formal. The family Scarabaeidae is considered here sensu lato to include
Geotrupidae. The described forms apparently belong strictly to Scara-
baeidae, but it is not always possible to substantiate it.

Genus Geotrupoides Handlirsch, 1906-1908

Type species. Geotrupes lithographicus Deichmiiller, 1886. Upper Jurassic
of western Europe.

Diagnosis. Rather broad, flat beetles. Head transverse, mandibles and
labrum clearly visible from under the clypeus. Pronotum transverse, laterally
rounded, rather long. Elytra with distinct punctate grooves.

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163

Species composition. One species in the Late Jurassic of western
Europe, two in the Early Cretaceous of Trans-Baikal.

Taxonomic position. The relations between Geotrupoides and other
Scarabaeidae remain unclear. Its position remains ambiguous even after the
study of new material which is far better preserved than the single specimen
of the type species. The genus is considered by us as formal since species in
a natural genus cannot differ so markedly as the newly described species do
from the type species. Labrum and mandibles of Geotrupoides project far out
from under the clypeus. Among Pleurosticti, only Glaphyrinae have a freely
articulate labrum but they lack distinctly punctate elytral grooves. Among
Laparosticti, Geotrupoides most closely resembles representatives of small
primitive families closely related to Hybosorinae, but an extensive com-
parison is not possible since the structure of antennae and abdominal spiracles
could not be studied in any of the specimens.

Geotrupoides sulcatus Nikritin, sp. nov.
(Plate X, Photos 4, 5; Figure 65)

Species name coined from ‘sulcatus’ (Latin)—grooved.

Holotype. No. 1668/1785, PIN, almost entire impression of beetle,
Trans-Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river
downstream from the mouth of Baisa river (Baisa site). Lower Cretaceous,
Valanginian-Goteriv, Zazin series.

Material. Besides holotype, three impressions of almost entire beetles
from the same site, specimen Nos. 1668/1801, 1989/2970, 3064/867, one of
which is dismembered into individual sclerites.

Description. Body broadly oval. Head almost equal in length and width,
widest at eyes and at temples, with small emarginations in front of eyes,
clypeus broad, rounded laterally, anteriorly truncate. Labrum rather long,
anteriorly blunt. Mandibles apically with well-developed denticles, consid-
erably projecting from under the clypeus. Pronotum transverse, its length less
than half its width, almost not narrowing anteriorly, laterally somewhat
rounded. Prosternum very short, much shorter than highly transverse
forecoxae. Width of forecoxae 2.5 times the length, with distinct keel.
Mesosternum long, with distinct posterior extension. Mesopleura large,
mesepisterna and mesepimera approximately equal in length, pleural sutures
almost anteriorly directed. Middle coxae oblique, strongly transverse. Scutel-
lum semicircular. Width of metasternum 2.5 times its length, almost not
anteriorly narrowing, metepisterna very slightly broadening anteriorly.
Posterior margin of metasternum angularly projecting backward. Hind coxae
large, slightly broadening laterally, its raised part running along metasternum
as a narrow band, extending backward along midline. Abdomen tapering
from base, sternites almost equal in length, freely articulating and may be

121

165

telescoped one under the other. Legs short, femora not extending beyond
lateral body margins. Femora broadened, forefemora 2.5 times and others 2
times longer than wide. Middle and hind femora each bearing two rows of
long hairs. Tibiae slightly shorter than femora, hind tibiae longer than middle
and foretibiae. Foretibiae flat and broad, with three denticles on outer margin
in anterior half, middle denticle noticeably larger than the remaining. Middle
tibiae not broad, with two transverse keels bearing stiff hairs. Spurs of middle
tibiae equal to two basal tarsal segments in length. Hind tibiae broadening,
its apical width one-third its length, with two transverse keels weaker than
those of middle tibiae. Keels and apices of tibiae with rows of stiff bristles.
Spur of hind tibiae long, equal in length to three basal tarsal segments.
Foretarsi small, half as long as tibiae, basal segment equal to the following
two together, second to fourth segments equal in length, last equal to the three
preceding. Middle and hind tarsi scarcely shorter than tibiae, their basal
segments equal to the two following, last segment slightly shorter than first.
Elytra with sharp, deep grooves, with indistinct punctures, grooves unite to
form pre-apical concentric pattern. Elytra crushed before their apices, ap-
parently they are strongly bent under fully covering the up of the abdomen
from behind. Head and pronotum with fine, dense punctation, elytra without
distinct punctation, punctation on underside of body fine, only mesosternum
with scattered, distinct, rather coarse punctation.

Dimensions. Body length 5.5-6.0 mm, width 2.5—3.0 mm; width of head
1.2 mm; length of pronotum 1.1—1.2 mm; elytral length 3.64.2 mm, width
1.3-1.5 mm; length of abdomen 1.5—2.5 mm.

Comparison. Differs from type species in small size and short pronotum,
from others in broad and flat tibiae.

Geotrupoides leptoscelis Nikritin, sp. nov.
(Plate X, Photo 6; Figure 66)

Species name coined from ‘leptos’ (Greck)—slender, and ‘skelidos’
(Greek)—leg.

Holotype. No. 3064/936, PIN, impression of beetle without antennae
and part of legs. Trans-Baikal, Buryat ASSR, Eravnin region, left bank of
Vitim river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Holotype.

Description. Body oval. Head transverse, widest behind eyes. Pronotum
transverse, its length two-thirds its width, slightly roundedly tapering before

Fig. 65. Geotrupoides sulcatus, sp. nov.; a, b—holotype PIN No. 1668/1785:

a—dorsal view, b—ventral view; c—paratype PIN No. 1989/2970, individual
sclerites. Baisa, Lower Cretaceous.

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166

ee aD)

\ y Toya
a ey YY b

Fig. 66. Geotrupoides leptoscelis, sp. nov.; holotype PIN No. 3064/936: a—dor-

sal view, b—ventral view. Baisa, Lower Cretaceous.

middle, pronotal angles almost right-angled. Prosternum short, slightly
shorter than transverse forecoxae. Mesosternum long, longer than middle
coxae, its transverse suture indistinct. Mesopleura large, mesepisterna some-
what longer than mesepimera, pleural sutures oblique. Middle coxae shorter
than wide, almost transverse to sternum. Metasternal length one-third its
width, laterally rounded, its posterior margin angularly extending backward.
Hind coxae large, not broadening laterally, its raised part running as a band
along posterior margin of metasternum, extending backward along midline.
Abdominal sternites almost equal in length, freely articulate. Legs rather
long, femora noticeably extending beyond lateral body margins. Femora
slightly uniformly broadening, hind femora slightly longer than middle
femora. Middle and hind tibiae equal to femora, slender, not broadening, with
longitudinal groove and one short transverse keel. Keel on the middle tibiae
beyond basal third, that on hind in the middle. Tibial spurs long and slender,
longer than basal tarsal segments. Elytra with sharp, slender grooves with
faint punctures. Body with fine, dense shagreened punctation, mesosternum
with additional large, well spaced punctures.

Dimensions. Body length 6.5 mm, width 3.0 mm; length of head 0.9
mm; length of pronotum 1.3 mm; elytral length 4.2 mm; width 1.5 mm; length
of abdomen 2.3 mm.

Comparison. Differs for the type species in smaller size, and pronotum
tapering toward apex, differs from other in slender legs with long tibiae and
narrow grooves on the elytra.

167

Geotrupoides vitimensis Nikritin, sp. nov.
(Plate X, Photo 7; Figure 67)

Holotype. No. 1668/1805, PIN, impression of almost entire beetle. Trans-
Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river downstream
from the mouth of Baisa river (Baisa site). Lower Cretaceous, Valanginian-
Goteriv, Zazin series.

Material. Holotype.

Description. Body broadly oval, Head transverse, widest near eyes.
Labrum large, wide, anteriorly blunt. Mandibles with apical denticles,
projecting from under the labrum. Length of pronotum two-thirds its width,
pronotum widest in anterior third, markedly narrowing from this point
anteriorly and towards base, laterally rounded. Prosternum very short, shorter
than forecoxae. Mesosternum shorter than middle coxae, mesepisterna
longer than mesepimera, middle coxae almost contiguous, oval, only slightly
shorter than their width. Scutellum triangular, apically acutely pointed.
Metasternum one-third its width, slightly narrowing anteriorly, posterior
margin angularly projecting. Raised part of hind coxae gradually reduced in
length laterally. Abdomen shorter than meso- and metathorax together, its
sternites short, equal in length. Legs short, femora only extending up to lateral
body margins. Femora rather strongly uniformly thickened; length of fore-
and hind femora two times, and that of middle femora 2.5 times the width.
Middle and hind femora each with two oblique rows of long hairs. Tibiae

pale SN
I"

a

Fig. 67. Geotrupoides vitimensis, sp. nov.; holotype PIN No. 1668/1805: a—dor-
sal view, b—ventral view. Baisa, Lower Cretaceous.

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168

approximately equal to femora in length. Foretibiae curved, flat, strongly
broadening anteriorly, in anterior third with three large deticles on outer
margin, and close to middle with short blunt dencticle. Middle and hind tibiae
slender, apically broadening, with transverse keel close to the middle, outer
margin opposite to keel projecting as broad blunt denticle. Basal segments
of foretarsus short. Elytra with sharp, deep, rather broad grooves, with dis-
tinct punctures, grooves uniting preapically in pairs to form concentric
pattern. Body dorsally almost smooth, vertrally with distinct scattered punc-
tures, middle coxae with slightly finer punctation.

Dimensions. Body length 8.7 mm, width 4.5 mm; length of head 1.2
mm; length of pronotum 1.5 mm; length of elytra 6.2 mm, width 2.2 mm;
length of abdomen 2.4 mm.

Comparison. Resembles type species in the shape of pronotum, but
differs from it in smaller size, differs from other species in the shape of
pronotum and structure of legs.

Genus Proteroscarabeus Grabau, 1923

Type species. P. yeni Grabau 1923. Lower Cretaceous of China.

Diagnosis. Large and medium-sized, broad, flattened beetles. Labrum
and mandibles not fully concealed under clypeus. Pronotum transverse,
laterally rounded. Spiracles of abdominal segments large, present on
membrane*. Elytra not covering tip of abdomen, without distinct punctate
grooves. Wings with two veins between Cubitus and the basally joining Anal.

Species composition. Type species P. yeni Grabau in the Early Creta-
ceous of China (Laiyan) and Trans-Baikal, second species in the Early
Cretaceous of Trans-Baikal.

Taxonomic position. Judging from the structure of hind wings, shape of
labrum, and arrangement of abdominal spiracles in membrane*, this form
must be related to the lower Laparosticti, including Geotrupinae; hind wing
most closely resembles wing of Geotrupes; a more accurate determination
of its taxonomic position is difficult owing to imperfect preservation. It is
probably an independent taxon of the same rank as the minor families of
primitive Laparosticti.

Proteroscarabeus baissensis Nikritin, sp. nov.
(Plate XI, Photos 1-4; Figure 68)

Species named after Baisa site.
Holotype. No. 1668/1830, PIN, impression of almost entire beetle.
Trans-Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river

*More correctly on intersegmental membranes—General Editor.

126

169

downstream from the mouth of Baisa river (Baisa site). Lower Cretaceous,
Valanginian-Goteriv, Zazin series.

Material. Besides holotype, isolated metathorax specimen No. 1989/
2885, two incomplete hind wings specimen Nos. 1668/1761 and 3064/1072
from the same site, two isolated elytra specimen Nos. 2385/1 and 2385/5
from the Pad Semen site, Trans-Baikal, Chita oblast, Chita region, Elizabetin
depression, Olengui river basin, Pad Semen, Lower Cretaceous, Argun
series. ;

Description. Body large, broadly oval, rather thick. Head small, much
narrower than pronotum, anteriorly almost not narrowing. Clypeus short,
transverse, anterior border margined, medially with small tunercle. Labrum
large, notably projecting from under the clypeus, anteriorly truncate, laterally
rounded, mandibles strongly projecting anteriorly and laterally. Length of
pronotum approximately two-thirds its width, widest in front of posterior
angles whence anteriorly tapering roundly; its anterior margin emarginate,
posterior margin strongly angularly extending backward, apically and at
posterior angles rounded. Backward curved transverse ridge running in
anterior third of pronotum. Prosternum short, forecoxae transverse. Sctutel-
lum large, wide, transverse, rounded apically. Mesosternum transverse,
noticeably shorter than middle coxae. Middle coxae convergent, ovoid,
slightly oblique, almost equal in length and width. Metanotum rather long,
half its width, alar cristae very sharp, scutoscutellar sutures indistinct, an-
tenotal membranous portions absent. Metasternum very short, less than
one-third its width, noticeably narrowing anteriorly, length of middle coxae
2.5 times the distance between middle and hind coxae. Metepisterna
noticeably broadening anteriorly. Hind coxae large, slightly broadening
laterally, their raised part in medial third projecting backward, its length
slightly less than width, raised part gradually tapered laterally, postero-lateral
angles of medial backward projection of coxa acute, extending. Abdomen
longer than meso- and metathorax together, not fully covered by elytra.
Abdominal sternites approximately equal in length. Abdominal spiracles
large, length of peritremes fully one-third to one-fourth the length of ab-
dominal sternites. Spiracles not symmetrical in relation to body axis, being
inamembranous fold. Legs rather long, forefemora not extending but middle
and hind femora noticeably extending beyond lateral body margins.
Foretibiae equal in length to femora, curved, flat and broad, with two
longitudinal rows of hairs, their pre-apical width one-fourth the length. Three
large denticles on outer margin in anterior half of tibiae, middle denticle
somewhat larger than remaining. Hind tibiae slightly shorter than femora,
flat, broadening strongly and uniformly from base to apex, with longitudinal
grooves, its lateral margin beyond middle and pre-apically with blunt den-
ticles. Spur of hind tibiae stiff, large, fully two-thirds the length of tibia itself.
Foretarsi equal in length to tibiae, attached subterminally in a pit on upper

7A

side of tibia, first segment slightly longer than the equal second and third,
fourth longer than first, fifth longer than fourth.

Elytra convex, strongly narrowing posteriorly, apically rounded.
Numerous longitudinal rows of faint punctures on elytra. Lateral third of
elytra covered with pubescence, two additional narrow pubescent long-
itudinal bands running along elytral suture and along middle. Hind wings not
very long, apical folded region approximately half the proximal region.
Anterior marginal vein distinct, with spiral thickening: large sclerotized
portion under it more proximal to the fold. R in the bent part sclerotized,
wide, broadening to flexure of wing. RS proceeding rather sharply backward,
distally curved along longitudinal axis of wing. M (reverse vein) reaching
almost to the wing base. Two veins not joining basally with other veins
present between Cubitus and first basally joining Anal vein. They are rather
stiff, particularly the anterior vein, the latter not differing in thickness from
other veins and basally passing close to the wing base. Surface of wing
between veins corrugated in areas close to the margin. Body without distinct
punctures dorsally, rough, somewhat tubercular. Metasternum along mar-
gins, especially in posterior corners, with large scattered tubercles; posterior
margin of the raised part of hind coxae with sharp, transverse indentations,
surface of coxae with dense distinct punctures.

Dimensions. Body length 35 mm, width 16 mm; length of head 6 mm;
length of pronotum 11 mm; length of elytra 16-17 mm, width 7-8 mm; length
of wing 26 mm; length of abdomen 13 mm.

Comparison. Differs from type species in slightly larger size, and more
strongly broadening hind tibiae.

Proteroscarabeus yeni Grabau, 1923
(Plate XI, Photos 5, 6; Figure 69)

Material. The stray remains of beetles resembling the holotype P. yeni in
structure and differing from the species described above in smaller size are
tentatively included in this species. These fragments include impressions of
a beetle lacking a head, pronotum and a large part of the legs, specimen No.
2385/2 from the Pad Semen site, Trans-Baikal, Chita oblast, Chita region,
Elizabetin depression, Olengui river basin, Pad Semen, Lower Cretaceous,
Argun series. Two impressions of forelegs, specimen Nos. 1668/1797
and1989/2638 and three impressions of elytra, specimen Nos. 1989/2946,
1989/2957 and 1989/2977 from Baisa site, Trans-Baikal, Buryat ASSR,
Eravnin region, left bank of Vitim river downstream from the mouth of Baisa
river, Lower Cretaceous, Zazin series.

Fig. 68. Proteroscarabeus baissensis, sp. nov.; a—holotype PIN No. 1668/1830;
b—paratype PIN No. 1989/2885, c—paratype PIN No. 1668/1761, d—paratype
PIN No. 3064/1072; Baisa, Lower Cretaceous, e—reconstruction of hind wing.

172

127 Fig. 69. Proteroscarabeus yeni Grabau; a—specimen PIN No. 2385/2; Pad Semen,

127

Lower Cretaceous; b—specimen PIN No. 1989/2638, Baisa, Lower Cretaceous.

Description. Forefemora broad, less than twice as long as wide, tibiae
shorter than femora, curved, broadening from base to apex. Outer margin of
tibia with tuft of suff hairs and in distal half with long denticles, two subequal
apical denticles much larger than following denticles. Tarsi attached subter-
minally to upper surface of tibia in a pit lateral to short curved spur. Tarsi
slender, basal four segments almost equal, shorter than following segment,
each with a pair of apical bristles; claws straight, weak, slender.

Middle and hind tibiae slender, almost not broadening toward apex, with
spines on each side. Abdominal sternites almost equal in length. Abdominal
spiracles large, peritremes one-third to one-fourth the length of sternites.

Dimensions. Body length of studied specimens about 20 mm, some-
what larger than indicated for the holotype; elytral length 12-14 mm, width
6 mm; length of abdomen 11 mm.

Comparison. Distinguished by spines along lateral margins of middle
and hind tibiae.

Genus Holcorobeus Nikritin, gen. nov.

Genus name coined from ‘holkos’ (Greek)—band, and genus Scarabaeus.
Type species. H. vittatus, sp. nov. Lower Cretaceous of Trans-Baikal.
Diagnosis. Small, broad, flat beetles. Head rather long, mandibles and

labrum strongly projecting from under the clypeus. Pronotum transverse,

short, laterally rounded. Elytra with indistinct punctate grooves and dark
longitudinal bands, not covering tip of abdomen. Wings with large folded
apical part, constituting almost half the length of the wings. RS parallel to

128

129

173

anterior margin of wing, almost entire folded part without veins, only with
corrugation. Claws without denticles.

Species composition. Two species in the Early Cretaceous of Trans-
Baikal.

Taxonomic position. In external appearance more closely resembles
May beetles than dung beetles. But, among the latter, only in Glaphyrinae
are the labrum and mandibles not covered by the clypeus. Holcorobeus is
evidently not affiliated to this subfamily since it has terminal spurs on hind
tibiae. The structure of antennae and abdominal spiracles is also not known
in Holcorobeus. Therefore its taxonomic position cannot be precisely deter-
mined, and the proposed genus should be regarded as formal.

Holcorobeus vittatus Nikritin, sp. nov.
(Plate XI, Photo 7; Figure 70)

Species name coined from vittatus (Latin)—banded.

Holotype. No. 3064/939, PIN, impression of almost entire beetle
without antennae. Trans-Baikal, Buryat ASSR, Eravnin region, left bank of
Vitim river downstream from the mouth of Baisa river (Baisa site). Lower
Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Besides holotype, impression of beetle without prothorax,
head and legs, specimen No. 1989/2961 and impression of isolated elytron,
specimen No. 1668/1809 from the same site.

Description. Body flat, broadly oval. Head slightly shorter than its
width, widest at occiput, with deep emarginations in front of eyes. Labrum
long, semicircular, scarcely less than clypeus in length. Clypeus anteriorly
with small border, medially narrowly emarginate, with tubercle at posterior
margin. Width of pronotum 2.5 times its length, widest close to middle,
uniformly narrowing anteriorly and posteriorly. Prosternum very short, much
shorter than transverse forecoxae. Anterior angles of pronotum acute, slightly
drawn out anteriorly, hind angles obtuse, rounded. Scutellum narrow, trian-
gular, pointed apically. Mesosternum small, shorter than middle coxae,
postepisterna of mesothorax oblique, extending along inner margins of coxae
anteriorly. Mesopleura large, mesepisterna and mesepimera equal in length,
mesepisterna medially almost reaching the middle coxae. Middle coxae
oblique, oval, apically contiguous, slightly longer than wide. Length of
metasternum less than width at posterior margin strongly tapering anteriorly,
posterior margin strongly angularly projecting backward, metepisterna
broadening anteriorly. Abdomen shorter than meso- and metasternum
together, narrowing almost from base, sternites equal in length, two-fifths of
large semicircular pygidum. Legs rather short, femoral apices reaching
lateral body margins. Femora uniformly broadening, forefemora two-thirds
the middle and hind femora. Foretibiae noticeably shorter than forefemora,

128

174

Fig. 70. Holcorobeus vittatus, sp. nov.; holotype PIN No. 3064/939: a—dorsal view,
b—ventral view, c—middle tibiae and tarsi, d—hind tibiae and tarsi. Baisa, Lower
Cretaceous.

flat, anteriorly broadening, with three denticles on outer margin in anterior
half; middle denticle much longer than remaining, with parallel sides and
rounded apex. Middle tibiae equal to foretibiae in length, approximately half
of femora, flat, broadening toward apex. Oblique transverse keel with tuft
of stiff short bristles running along middle of tibia, similar tuft also present
along apex. Spurs of middle tibiae scarcely longer than first tarsal segment,
scarcely differing in thickness. Hind tibiae equal in length to femora, flat,
strongly broadening from base to apex, divided into equal parts by two

ers

130

175

oblique transverse keels bearing tuft of stiff short bristles as at the tibial apex.
Spurs of hind tibiae narrow, scarcely longer than first tarsal segment. Middle
tarsi noticeably longer than tibiae, their segments slightly broadening toward
apex, first tarsal segment equal to last, more than twice as long as second,
second to fourth equal in length. Claws slightly curved, weak, uniform,
slightly shorter than fourth tarsal segment. *Scarcely longer than tibia,
segments noticeably broadening toward apices, first segment equal to fifth,
one-and-a-half times longer than second, second-fourth equal. Claws
markedly curved, weak, uniform, two-thirds of fourth tarsal segment. Elytra
fully covering the body laterally, scarcely covering base of pygidium, bearing
12 broad, shallow grooves, with indistinct large punctures. Body with fine,
dense punctation, punctures somewhat sharper on mesosternum and
pygidium. Pronotum with dark-colored border, scutellum with two lon-
gitudinal dark-colored bands, elytra with dark-colored pattern consisting of
broad bands along suture, bands commencing from shoulder obliquely
running to sutural band and joining with it in apical third of elytron, and dark
apical spot on outer margin narrow; dark-colored lines running along all
punctate furrows.

Dimensions. Body length 10.5—12.0 mm, width 5.5—6.5 mm; length of
head 1.9 mm; length of pronotum 1.6 mm; length of elytra 6.5—7.5 mm, width
2.9-3.2; length of abdomen 3.5 mm.

Comparison. Differs from second species in structure of tarsus and
pattern on elytra, in which dark-colored bands are joined pre-apically.

Holcorobeus picturatus Nikritin, sp. nov.
(Plate XI, Photo 8; Figure 71)

Species name coined from ‘picturatus’ (Latin)—painted.

Holotype. No. 1989/2994, PIN, impression of beetle without head and
prothorax, with unfolded elytra and hind wings. Trans-Baikal, Buryat ASSR,
Eravnin region, left bank of Vitim river downstream from the mouth of Baisa
river (Baisa site). Lower Cretaceous, Valanginian-Goteriv, Zazin series.

Material. Besides holotype, impression of almost entire poorly pre-
served beetle, specimen No. 1989/2926, and isolated elytron, specimen No.
3064/850 from the same site.

Description. Body flat, oval. Length of head markedly less than width,
in front of eyes with shallow narrow emarginations. Length of pronotum
two-fifths its width. Forecoxae transverse, with blunt keel. Scutellum trian-
gular, slightly longer than wide. Middle coxae elongate along body, con-
tiguous to a large extent apically, slightly longer than wide. Length of

*Translation of the following sentence correct. Apparently the words ‘Hind tarsi’ missing in
the Russian original. Otherwise the sentence does not make any sense—General Editor.

176

metasternum one-third the width at posterior margin, strongly narrowing
anteriorly, posterior margin slightly extending backward. Abdomen shorter
than meso- and metathorax together, tapering from base of third sternite.
Femoral apices noticeably extending beyond lateral body margins. Femora
broadening, fusiform. Foretibiae shorter than femora, flat, with a rather large
blunt denticle in anterior third, more proximal to it and apically with small
denticles. Middle tibiae short, much shorter than femora, flat, broadening
toward apex. Oblique keel with tuft of short stiff bristles present in distal
third; similar tuft but with much larger bristles at broad apex of tibia. Spurs
of middle tibiae narrow, slightly longer than first tarsal segment. Hind tibiae
shorter than femora, broadening in apical third with transverse oblique keel
and with tuft of stiff hairs as at apex. Spurs of hind tibiae large, flat, cultrate,
fully half the length of tibiae, slightly longer than first tarsal segment. Middle
tarsi scarcely longer than tibiae, their segments noticeably broadening toward
apex, first segment three times as long as second, second to fourth segments
equal, last two-thirds of first. Claws uniform, curved, equal to fourth tarsal
segment. Hind tarsi longer than tibiae. Elytra without noticeable grooves.
Body with fine punctation. Elytra with dark-colored, posteriorly narrowing
band along suture and slightly oblique band running from shoulders to apex.

Dimensions. Body length 9.5-10.5 mm, width 4.5 mm; length of head
1.8 mm; pronotum 1.2 mm; elytra 6.6 mm, width 2.8 mm; length of wing
12.0 mm; length of abdomen 4 mm.

Comparison. Distinguished by structure of tibiae and tarsi and elytral
pattern consisting of parallel dark-colored bands.

Fig. 71. Holcorobeus picturatus, sp. nov.; holotype PIN No. 1989/2994.
Baisa, Lower Cretaceous.

nag cab

13

—

177

INFRAORDER ELATERIFORMIA

SUPERFAMILY CEBRIONOIDEA LATREILLE, 1802
(= Elateroidea Leach, 1815)!

Family CEROPHYTIDAE Latreille, 1834

This family includes a single extant genus Cerophytum Latr. with several
species from western Europe, North and Central America. Fossil repre-
sentatives of Cerophytidae were not known until now. As it turns out, remains
of these beetles are frequently encountered in the Cretaceous fossil resins of
Siberia.

Based on several distinguishing characters, the Cretaceous genus
Aphytocerus described below may be considered less advanced than the
extant Cerophytum. In Aphytocerus the antennae are only slightly serrate
even in the male (i.e., the process of increasing the surface of their segments
had only just commenced), whereas Cerophytum males possess highly
serrate antennae. The development of serrate antennae is usually associated
with complicated sensory apparatus. Further, in Aphytocerus the femoral
plates still exist, although greatly reduced in comparison with many other
Cebrionoidea. They are completely absent in Cerophytum, which Crowson
(1955) explains by the necessity to increase the surface area of the femora in
connection with the development of the springing ability. In Aphytocerus the
hind femora do not exhibit any characters for this purpose. The slightly
oblique articulation of the trochanters with the femora, distinguishing
Cerophytum from other Cerbrionoidea, is probably a secondary phase which
Aphytocerus also has not reached. Moreover, many apomorphs common to
Cerophytum and Aphytocerus (marked elongation of trochanters, develop-
ment of antennal sockets, flattening of hind coxae, and others), must have
already been formed before the Late Cretaceous. However, Aphytocerus has
several autapomorphs which prevent its consideration as an ancestor of
Cerophytum (head deeply retracted into prothorax, spinescent prosternal
process). Aphytocerus is undoubtedly an extinct specialized branch of the
family Cerophytidae.

Some postulations can be made on the ecological characteristics of
Aphytocerus and the direction of its specialization. As far as is known, species
of the genus Cerophytum in Europe and North America live under the bark
of dead trees and in decomposing wood. This is probably the original mode
of life for Cerophytidae. In its general habits Aphytocerus somewhat
resembles Anobiidae, living in dry, dead wood and preferring dead knots on
living trees. Inhabitants of decomposed wood are rather uncommon in fossil
resins. They apparently occur by chance, since repeated finds of any par-

‘The name Elateroidea Leach, 1815, suggested by Leng (1920) must be replaced by the

available priority name Cebrionoidea Latreille, 1802, nom. trans. hic ex Cebrionides Latreille,

1802.

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ticular species are rare. Species inhabiting living trees are far more common
in resin. Aphytocerus is observed in resin very frequently. It is present in a
small volume of material of the Senomanian resins from the Ust’-Yenisey
basin, while in the collection of Cognacsantonian resin from the Khatang
basin it is represented by seven specimens, constituting 25% of the total
number of beetles. Therefore it is highly likely that Aphytocerus inhabited
the same species (or several species) of conifers which produced the resins
containing their remains. Unfortunately, there is still no information on the
taxonomic status of these conifers; judging from the composition of the
spore-pollen conglomerates, Cupressaceae and/or Taxodiaceae as well as
Pinaceae and Pinus in particular were predominant among conifers in Taimyr
at that time (Saks et al., 1959). The development of Aphytocerus probably
took place in dead knots of trees at the base of which resin secretion is
frequently observed in the extant conifers; this mode of life was definitely
most suited for burial.

Genus Aphytocerus Zherichin, gen. nov.

Genus name coined from genus Cerophytum.

Type species. Aphytocerus communis, sp. nov. Upper Cretaceous of
Taimyr.

Description. In general body shape this species resembles repre-
sentatives of the family Anobiidae; covered by sparse short hairs.

Head markedly retracted into prothorax and bent under it, dorsally
almost not visible. Eyes large, rounded, occupying large part of visible
surface of head. Antennae ten-segmented, attached to frons between eyes in
deep, wide, rounded sockets; their inner margins continuous while outer
margins directly bordering the eyes. Distance between antennal bases almost
equal to the length of their first segment. Antennae fairly long, extending
beyond base of elytra, weakly serrate; their first segment very large, apically
truncate, last segment slender, longer than penultimate. Antennae slightly
shorter in males than in females and more strongly serrate. Maxillary palps
short with dolabriform broadening of last segment. Mouth parts at rest
covered by anterior hood of pronotum and not visible.

Protonum transverse, companulate, highly convex, rounded laterally.
Convexity of pronotum not same all over; highest in anterior half, pronotum
hanging somewhat like a hood over the head. Posterior angles of pronotum
drawn out into short cusps, its base with two emarginations. Pronotum at
anterior margin with distinct hood, with pronounced, rather deep, arcuate
longitudinal grooves for insertion of antennae. Posterior margin of proster-
num extending into narrow point, inserted into mesosternal depression and
almost reaching midpoint of middle coxae. Scutellum large, wide, flat,
apically rounded.

WW)

Elytra elongate, almost cylindrical, with angularly projecting shoulders
and distinct humeral tubercle, apically broadely rounded, fully covering
abdomen. Basal margin of elytra arcuate, rounded. Lateral margins almost
parallel, in side view with sharp S-shaped bend at the level of hind coxae.
Punctate grooves regular, with distinct and rather deep punctures; last groove
truncate, reaching only up to hind coxae.

Mesosternum very short, with depression for insertion of prosternal
process. Metasternum large, slightly convex, with small medial groove at
posterior margin.

First abdominal sternite short, following three of same length, slightly
longer than first and weakly convex. Fifth sternite nearly as long as fourth,
flat, apically rounded, without pits or elevations. Sutures between sternites
straight.

Forecoxae rounded, convergent, separated only by narrow prosternal
process. Middle coxae the same size, spherical, protuberant, convergent,
distance between them about one-third of their diameter. Hind coxae strongly
transverse, almost completely flat, with small femoral plates. Trochanters
very large, obliquely articulate with femora, their length in fore- and middle
legs about one-third and in hind legs one-half of the femoral length. Femora
weakly clavate, with subapical constriction and notch for articulation;
forefemora* almost round in cross section, slender, in length nearly equal to
femora plus trochanters, in apical half covered by rather long (only slightly
shorter than tibial diameter), slender, vestigial hairs, with two slender, short
apical spurs on inner side. First tarsal segment cylindrical, long; second and
third equal in length, half the length of first, more or less triangular; fourth
somewhat longer and much wider than third, flattened; fifth long and slender,
thickening apically, attached in depression on inner side of fourth segment.
Foretarsi slightly broadening in male. Claws slender, long almost half the
length of tarsal segment, slightly curved, with weak, blunt basal denticle.

Species composition. Two species in the Late Cretaceous of Taimyr.

Comparison. Differs from sole earlier known genus Cerophytum in
serrate (not pectinate) antennae of male, presence of femoral plates (though
reduced) on hind coxae, oblique articulation of trochanters with femora, head
deeply retracted into prothorax, and narrow and angular (bit not rounded)
prosternal intercoxal process.

Remarks. The combination of characters such as serrate ten-segmented
antennae, rounded forecoxae, grooves for insertion of antennae, posterior
prosternal process, strongly transverse and almost contiguous hind coxae
with femoral plates, absence of transverse suture on metasternum, five-seg-
mented tarsi, and the trilobate male genitalia make it possible to confidently
include Aphytocerus under Cebionoidea. Exceptionally long trochanters; flat

*An obvious misprint in the Russian original. Logically should refer to tibiae—General
Editor.

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180

hind coxae; elongate first and broad fourth tarsal segment; first segment of
antennae enlarged, inserted in large, closely convergent sockets between
eyes, and presence of small pronotal hood are characteristic in this super-
family only of Cerophytidae; the latter is a small independent group, sharply
differing from the other families in several characters. R. Crowson (1955),
in the diagnosis of the family Cerophytidae describes several atypical
charcters of Aphytocerus (such as slightly oblique articulation of trochanters
with femora and total reduction of femoral plates of hind coxae). Since this
diagnosis is based on a single extant genus, it is not surprising that the find
of a second genus necessitates a certain alteration in the diagnosis.

Aphytocerus communis Zherichin, sp. nov.
(Plate XII, Photo 1; Figure 72, a—e)

Species name coined from ‘communis’ (Latin)—common.

Holotype. No. 3311/41, PIN, male, a fully preserved beetle embedded
in fossil resin (retinite). Taimyr national district, Khatang region, Yantardakh
area on the right bank of Maimechi river 5 km upstream from its mouth
(Yantardakh site). Upper Cretaceous, Gognac-Santon, Kheta series.

c d
aS
e f

Fig. 72. Representatives of genus Aphytocerus: a to e—A. communis, sp. nov.;

holotype PIN No. 3311/41: a—head and prothorax, b—genitalia, c—antenna,

d—tarsus, e—hind tibia; Yantardakh, Upper Cretaceous; f—A. dolganicus sp. nov.;
holotype PIN No. 3426/35, hind tibia; Agapa, Upper Cretaceous.

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181

Material. Besides holotype, six more or less damaged and incomplete
remains from the same site, specimen Nos. 3130/28, 3130/29, 3130/35,
3311/42, 3311/43, 3311/44.

Description. Color of body black. Eyes, palps, antennae, apices of
forefemora, middle and hind trochanters and femora, all tibiae and tarsi light
yellowish-brown, first segment of antennae and middle of middle and hind
legs dark colored. Head, pronotum, elytra, and thorax covered with very
dense and minute, almost pollinose, fine hairs; abdominal sternites with
rather long and very slender, contiguous, whitish hairs.

Frons anterior to antennal sockets with fine, dense punctation. Antennae
in male extending beyond humeral tubercles of elytra, distinctly serrate,
dorso-ventrally flattened. Their first segment broad, oval, slightly longer than
wide; following five segments similar, triangular, almost equal in length and
width; seventh and eighth of the same length but narrower, with less project-
ing apical corners; ninth equal in length to eighth, oval; tenth noticeably
narrower and somewhat longer than ninth, slightly pointed apically. Anten-
nae of female more slender and longer, reaching the level of hind coxae. Their
segments narrower and more elongate, with less projecting corners, but also
flattened; length ratios of segments approximately the same as in the male.
All antennal segments densely covered with short and slender hairs.

Pronotum widest in middle, equal in width to elytra, width 1.25 times
the length, with small lateral depression in front of hind comers, depression
deeper in male, posterior angles directed backward, somewhat laterally and
upward. Sculpture formed by very dense, fine punctures, distance between
which almost equal to their diameter; pattern very fine-grained laterally.

Ratio between elytral length and width 14 : 9. Sides of elytra behind
anterior third with small but distinct depression. Punctures in grooves elon-
gate, distance between punctures considerably less than their length; inter-
vals between punctate grooves narrow, equal in width to grooves but laterally
narrower and very finely shagreened. Lower side of thorax, coxae, femora,
trochanters and abdominal sternites with very fine, dense, uniform punctures,
distance between which nearly equal to their diameter. All tibiae straight.
Genitalia as in Plate XII, Photo 1b; Figure 72b.

Dimensions. Body length 2.6—-3 mm.

Comparison. Distinguished by denser, elongate punctures in elytral
grooves and straight hind tibiae.

Aphytocerus dolganicus Zherichin, sp. nov.
(Figure 72f)

Species named after Dolgan nationality.
Holotype. No. 3426/35, PIN, male, split into two parts, and partially
oxidized inclusion of beetle in fossil resin (retinite), apex of elytra not

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182

preserved. Taimyr national district, Ust’-Yenisey region, Lower Agapariver,
40 km downstream from its mouth (Agapa site). Upper Cretaceous,
Senomanian, Dolgan series.

Material. Holotype.

Description. As far as can be judged from the preserved fragments of
integument, the body color is black. Antennae and legs (except coxae)
entirely light yellowish-brown. Pubescence as in A. communis.

Antennae in male reaching the level of hind coxae, distinctly serrate,
dorso-ventrally flattened. Their first segment broad, oval, longer than wide;
following four segments similar, triangular, somewhat longer than wide;
sixth, seventh and eighth segments the same length, slightly narrower, with
less projecting apical corners; ninth segment oval, equal in length to eighth;
tenth almost negligibly narrower and longer than ninth.

Pronotum widest at middle, equal to elytra in width, width 1.25 times
the length, laterally uniformly rounded, anterior to posterior angles with
shallow depression. Sculpture formed by very dense, fine punctures, distance
between punctures in the middle of pronotum slightly more than their
diameter but almost equal on sides near posterior margin.

Elytra, behind anterior third, with lateral depressions of apparently about
the same proportions as in A. communis. Punctures in grooves round, distance
between them 1.5—2 times their diameter. Intervals between punctate grooves
narrow, nearly one-and-a-half times broader than grooves, but laterally equal
to them in width, and finely shagreened. Lower side of thorax, coxae, femora,
trochanters and abdomen with very fine, dense, uniform punctures. Fore- and
middle tibiae straight, hind tibia distinctly curved in apical third. Structure
of male genitalia not known since, although preserved even in the holotype,
they are highly oxidised and the structural details are not discernible.

Dimensions. Body length 3.3 mm.

Comparison. Close to A. communis, differs from it only in larger size,
entirely light-colored legs, long antennae of male, round and sparser punc-
tures in elytral grooves, and curved hind tibiae.

SERIES OF SUPERFAMILIES OF CICUJIFORMIA

SUPERFAMILY CLEROIDEA LATREILLE, 1802
Family ACANTHOCNEMIDAE Crowson, 1964

This family was established for a single Recent genus Acanthocnemus Perr.
and until that time was unknown as a fossil. Initially it was given the rank of
a subfamily (Crowson, 1964), but later of a family (Crowson, 1970) based

_ ona study of larvae tentatively identified as Acanthocnemus. The mode of

life of Acanthocnemidae is almost unknown. Crowson (1970, p. 19) suggests

183

that they are affiliated to the main line of “more or less predatory” cleroids,
but pollen also forms an important food for several members of other families
in this branch. The young larva was found in soil under the roots of eucalyptus
and is apparently a predator.

Acanthocnemidae is morphologically one of the most archaic groups
among Cleroidea and probably one of the most ancient families of this
superfamily. The extant genus Acanthocnemus Perr., judging from its isola-
tion and unique geographic distribution, is a very ancient relict. According
to G.G. Yakobson (1905-1915) and G. Champion (1922) only one species
A. nigricans (Hope) belongs to this genus, and has been repeatedly described
under different names. The geographic distribution of this species covers
Australia, Tasmania, New Caledonia, Burma, Thailand, India, Madagascar,
South Guinea, Algeria, Sicily, Sardinia, Corsica and Cyprus. Champion
considers Australia to be its native habitat and suggests that it has been
introduced into other parts of the world. However, Acanthocnemus has
apparently never been noted in ships or in transported goods. It does not occur
solely in the Port areas, nor has it been found even once in the western
hemisphere. Therefore one may suspect that A. nigricans inhabits a very wide
but distinctly discontinuous natural geographic area; probably, it existed as
a species for a very long time, at least since the Early Paleogene. Examples
of such ancient species, though very few, do occur among insects. In some
instances, the habitats of present-day species of terrestrial arthropods
resemble, to some extent, the habitat of A. nigricans, for example, those of
the Protura (Tuxen, 1967).

Genus Acanthocnemoides Zherichin, gen. nov.

Genus name coined from Acanthocnemus Perr.

Type species. Acanthocnemoides sukatshevae, sp. nov. Upper Creta-
ceous of Taimyr.

Description. Body somewhat flat, resembles members of the Mala-
chiinae (Melyridae) in general appearance. Uniformly covered with
moderately dense, thick, erect bristles. Integument apparently rather soft, but
not to the same extent as in the Malachiinae.

Head broad, its width (with eyes) greater than length. Eyes large,
rounded, highly convex, extending far beyond the margin of head. Frons
broad, only slightly narrower than anterior margin of pronotum. Clypeus
simple, triangular. Temples long, nearly half the length of eyes. Mandibles
with two apical denticles. Maxillary palps with triangular segments broaden-
ing toward apex and last segment terminally truncate. Labial palps short, last
segment dolioform. Antennae slender and rather long, extending beyond base
of elytra, attached in front of eyes at a very short distance from them. All

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184

antennal segments longer than wide; last three segments flattened and
broadened, forming a very loose but distinct club.

Pronotum transverse, with strongly rounded sides and distinct, rather
broad (particularly in posterior half), flat lateral margin. Posterior margin of
pronotum slightly wider than anterior, straight. Disk of pronotum weakly
convex. Prosternum without a depression in front of forecoxae, only with
very indistinct transverse fold. Scutellum triangular, weakly transverse.

Elytra long, ovoid, basally slightly broader than base of pronotum,
smoothly broadening posteriorly, widest behind midlength, apically broadly
rounded. Shoulders projecting, rounded, humeral tubercle nearly un-
developed. Epipleura broad at base, posteriorly narrowing, disappearing at
level of hind coxae.

Mesosternum very short, convex, its narrow pointed intercoxal process
almost reaching posterior margin of coxae. Metasternum long, weakly con-
vex, without depression or grooves; at anterior margin between coxae with
broad, short and flat, apically rounded projection; posterior margin of
metasternum arcuate. Abdominal sternites almost flat, similar, last sternite
in both sexes without pit or processes, broadly rounded at apex.

Forecoxae large, projecting, strongly convergent, distance between
them almost equal to tibial* width. Forecoxal cavities broadly open
posteriorly. Middle coxae rounded, convergent, separated only by narrow
mesosternal process. Hind coxae strongly transverse, contiguous, without
femoral plates. Legs slender and long. Trochanters rather large, obliquely
articulating with bases of femora. Femora long, slightly thickened toward
apex. Tibiae slender, equal in length to femora, outer margin without spines,
inner margin with row of short and thick erect bristles; [tibiae] straight, in
females hind tibiae slightly S-shaped. Tarsi equal to tibiae in length, slender,
five-segmented, with long cylindrical segments and long, slender, simple
claws. Last tarsal segment 1.5 times longer than penultimate. Foretarsi of
same structure in male and female. Hind tarsi slightly longer than other tarsi.

Species composition. One species from the Cretaceous of Taimyr.

Comparison. Differs markedly from the single earlier known genus
(Acanthocnemus Perr.) in the triangular (not cylindrical) last segment of
maxillary palps; elongate (not transverse) fourth to eighth antennal segments;
long, posteriorly broadening temples; ovoid (not parallel-sided) elytra; ab-
sence of pit on prosternum and row of spines on outer margins of tibiae;
shorter last tarsal segment, and identical structure of foretarsi in males and
females.

Remarks. This genus definitely belongs to the superfamily Cleroidea.
This is confirmed by the general appearance of the beetle and by the
projecting forecoxae, lack of plates on hind coxae, and structure of antennae

*So given in the Russian original. Earlier the intercoxal distance was always compared with
coxal and not tibial size, hence the doubt—General Editor.

137

278

185

and tarsi. The identification of the family is a somewhat more complicated
problem. Crowson (1970) characterizes the Acanthocnemidae by the follow-
ing structural features of the imago, which differentiate them from the
Melyridae: antennae with distinct three-segmented club; claws simple; outer
margin of tibiae with short, thick spines; prosternum with paired pits in front
of coxae; and edeagus with divided phallobase*. Of these characters, the first
two are observed in Acanthocnemoides but not next two; the structure of the
phallobase could not be determined from the available material. However,
based on these two characters, Acanthocnemoides is closer to Melyridae.
Crowson seems to attach little importance to the prosternal pit since he states _
“the first acanthocnemoids were possibly without prosternal pits” (Crowson,
1970, p. 19). The armature of the tibiae also seems to be an unimportant
character, and hence Acanthocnemoides is here assigned to the Acanthoc-
nemidae with which its proximity cannot be doubted. The Acanthocnemidae
may be better ranked as a subfamily of Melyridae. However, this question
cannot be adequately resolved in the absence of material on Recent Acan-
thocnemus.

Acanthocnemoides sukatshevae Zherichin, sp. nov.
(Plate XII, Photo 2; Figure 73)

Species name after I.D. Sukacheva, who collected the material.

Holotype. No. 3308/1, PIN, female, inclusion of an entire beetle in
retinite (split during study); Taimyr national district, Khanag region, right
bank of Zhadnikha river at its source (Zhadnikha site). Cretaceous, Albian
?—Senomanian, Begichev series (middle member).

Material. Besides holotype, one specimen (male), specimen No. 3308/2
slightly damaged, from the same site; found in the same lump of retinite as
the holotype.

Description. General coloration of body dark, blackish-brown; palps,
antennae and legs light-brown, last two segments of antennae black. Entire
body uniformly densely covered with fine, rather long, dark-colored, op-
pressed pubescence.

Head with fine, uniform punctation. First antennal segment large,
strongly thickening apically. Second slightly narrower than first, but wider
than subsequent segments, short, ovoid, slightly compressed dorsoventrally.
Following three segments of the same shape and size, slightly broadening
toward apex, not flattened, twice as long as wide; sixth segment of the same
width but slightly shorter, 1.5 times longer than wide; seventh 1.5 times
longer and slightly wider than sixth; eighth of the same size and shape as the
sixth. All these segments, except the first two, with whorls of fine hairs on

* Although the Russian original uses the term “tegmen” we prefer to use the more current
term “phallobase’”—General Editor.

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186

9»

QS5min

a

Fig. 73. Acanthocnemoides sukatshevae, sp. nov.; holotype PIN No. 3308/1:
a—dorsal view; b—ventral view; c—antenna. Zhadnikha, Cretaceous.

the broadened apical part. Ninth and tenth segments twice as wide as eighth,
dolioform, longer than wide; eleventh a little wider and longer than tenth,
apically pointed. All segments of club with rather long, sparse hairs.

Length of pronotum two-thirds its maximum width, and with fine, dense,
uniform shagreening. Pubescence directed posteriorly slightly longer on
sides than on disk. Elytra twice as long as maximum width, uniformly finely
shagreened, without grooves; pubescence on elytra directed posteriorly.
Thorax on ventral side and abdomen with similar shagreening.

First three segments of foretarsi equal, cylindrical, three times longer
than wide; fourth segment of same length but considerably thinner; fifth
equal in length to the preceding two together. First four tarsal segments
ventrally with stiff bristles. Middle tarsi of same structure; hind tarsi with
slightly longer segments, their length ratios same as in fore- and middle tarsi.

Only narrow apex of edeagus visible in paratype (male).

Dimensions. Body length 2.2 mm.

187
INFRAORDER CUCUJIFORMIA LATREILLE, 1802

Family CRYPTOPHAGIDAE Frichson, 1845
Subfamily Atomariinae Leconte, 1862
Genus Nganasania Zherichin, gen. nov.

Genus named after the Nganasan nationality.

Type species. Nganasania khetica, sp. nov. Upper Cretaceous of
Taimyr.

Description. Body slightly flattened dorsoventrally, with fine hairs.
Head considerably narrower than pronotum, transverse. Eyes large, rounded,
moderately convex, projecting beyond margin of head. Temples absent.
Antennae freely [= movably] attached to anterior part of frons in front of
eyes, rather long and thick, reaching humeral tubercles of elytra; basal
segments of antennae thickened, first segment longer and broader than
- second; antennal club loose, three-segmented.

Pronotum weakly transverse, in lateral view appearing to form a com-
mon flat arc with elytra, not delineated from the latter by a drop-off. Sides
of pronotum almost parallel in posterior half, converging anteriorly, without
denticles of projections. Widest part of pronotum is its weakly bicrenate base.
Anterior angles of pronotum not projecting, without depression; posterior
angles slightly projecting laterally; pronotum lacking discrete lateral margin.
Disk of pronotum with smooth ornamentation, without keel or depression.
Forecoxal cavities open posteriorly. Scutellum large, clearly visible, equal
in length and width, flat, pentagonal.

Elytra almost parallel-sided, basally slightly wider than pronotum, with
slightly projecting shoulders, posteriorly bluntly tapering, fully covering
abdomen. Sculpturing on elytra consisting of small punctures, not forming
distinct rows; presutural row of punctures apparently rudimentary but
impossible to confirm in our material. Epipleura of elytra developed in basal
half, rather wide, disappearing further on.

Mesothorax short, sloping backward, but not vertically; metathorax
long. Abdominal sternites weakly convex; third and fourth sternites of same
length; structure of first and second not known; fifth flat, rounded at apex.

Forecoxae transverse, projecting, closely convergent. Middle coxae
strongly convergent, rounded. Hind coxae small, triangular, closely conver-
gent. All trochanters large. Femora moderately thickened. Tibiae slender,
straight, equal in length to femora; foretibiae apically with two slender
indistinct spurs. All tarsi five-segmented (? female), their first four segments
identical in shape, simple, not bilobed; fifth segment long, equal in length to
the preceding three together; claws simple, slender and short.

Species composition. Monotypic genus.

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188

Comparison. Apparently very close to the extant genus Ootypus
Ganglb., but differs in pubescent integument. Also resembles Atomaria
Steph. but differs in the lack of a drop-off between the pronotum and elytra,
and by the pronotum not narrowing posteriorly.

Remarks. The affiliation of Nganasania to Cryptophagidae has been
reliably established from a complex of distinguishing characters (structure
of antennae and tarsi; converging coxae; posteriorly open forecoxal cavities;
long trochanters; presence of two spurs on foretibiae; epipleura of elytra dis-
appearing in posterior half, and to the subfamily Atomariinae from the
antennae freely [= movably] attached to the anterior part of frons. In its
general evolutionary level, Nganasania does not differ from the extant
Atomariinae, nor does it show any plesiomorphic features. The present-day
representatives of this group occupy various cryptic habitats ({forest] litter,
decomposing wood, burrows and nests of animals); they feed primarily on
mold fungi, rarely on plant shoots.

Neganasania khetica Zherichin, sp. nov.
(Plate XII, Photo 3; Figure 74)

Species name coined from Kheta series.

Holotype. No. 3311/45, PIN, inclusion of an entire beetle in fossil resin
(retinite); partially hidden by detritus and opacities in the resin. Therefore, it
was not possible to identify some structural details (particularly of the ventral
side). Sex could not be determined. Taimyr national district, Khatanga
region, Yantardakh area on right bank of Maimechi river, 5 km upstream
from its mouth (Yantardakh site). Upper Cretaceous, ? Kon’ yak-Santonian
Kheta series.

Ane

Q5mm
: d

Fig. 74. Nganasania khetica, sp. nov.; holotype PIN No. 3311/45: a—dorsal view;
b—ventral view; c—foreleg; d—antenna, Yantardakh. Upper Cretaceous.

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189

Material. Holotype.

Description. Coloration of body rusty-brown, epipleura of elytra slight-
ly lighter in color; pubescence short and fine, dark-colored. Bases of antennae
widely spaced, separated from each other by length of their first segment,
and from anterior margin of eyes by distance equal to half the diameter of
eyes. First antennal segment twice as long as wide; second segment slightly
thinner and three-fourths the length of first; third segment half as thick as
second, considerably longer than wide; following four segments alike, more
or less cylindrical, of same thickness but considerably shorter than third,
equal in length and width. Eighth segment of same length and width as
preceding segments, almost spherical. Ninth twice as wide as eighth, equal
in length and width almost square; tenth slightly wider than ninth, equal in
length and width, slightly broadening apically. Eleventh of same width as
tenth, slightly longer, symmetrical, pointed apically.

Pronotum one and a quarter times wider than long, and weakly
shagreened. Elytra three times as long as pronotum, with fine punctures and
weak shagreening. Lower side of body with similar ornamentation. Last
abdominal sternite with longitudinal groove. First four segments of tarsi
slightly longer than wide.

Dimensions. Body length 2.0 mm.

Family LATHRIDIIDAE Redtenbacher, 1845

Subfamily Corticariinae Stein, 1877
Genus Succinimontia Zherichin, gen. nov.

Genus name coined from ‘succinum’ (Latin)—amber, and ‘mons’ (Latin) —
mountain (Latin translation of the locality name Yantardakh).

Type species. Succinimontia infleta, sp. nov. Upper Cretaceous of
Taimyr.

Description. Body elongate, slightly flattened. Head broad, transverse,
narrowing anteriorly, together with eyes narrower than anterior margin of
pronotum. Eyes rounded, strongly convex, almost hemispherical. Temples
absent. Frons flat, without keels or tubercles. Fronto-clypeal suture distinct,
but shallow. Clypeus and frons coplanar. Antennae 11-segmented; third to
fifth segments round, equal in length and breadth; sixth to eighth segments
weakly transverse; club loose, three-segmented, its apical segment symmetri-
cal, pointed.

Pronotum transverse, wider in anterior half, posteriorly narrowing, with
weak denticles at sides. Posterior margin of pronotum slightly arched,
convex. Disk of pronotum without keels or longitudinal depressions; basal
pit absent, only a very flat [shallow] depression present, occupying entire

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base. Scutellum clearly visible, comparatively narrow, equal in length and
width, without pits, grooves, keels or folds; flat.

Elytra noticeably broader than pronotum, with rounded sides, ovoid,
broadest beyond middle, rounded apically, with rows of punctures. Intervals
between grooves without keels or projections. Development of pubescence
on elytral integument unclear; if present, then composed of poorly noticeable
short hairs. Humeral tubercles distinct, projecting, depression present be-
tween them and lateral margin.

Forecoxal cavities posteriorly closed. Abdomen with five visible ster-
nites. Projection of first sternite between hind coxae wide, flat, arcuate. First
sternite long, equal in length to the following two together, without femoral
lines. Second and third sternites equal in length, fourth slightly shorter;
sutures between sternites straight. Fifth sternite longer than fourth, apically
rounded, flat, without depressions or tubercles.

Forecoxae contiguous, middle and hind coxae wide-set. Trochanters
short, almost equal in length and width, obliquely articulating with femora.
Femora weakly clavate, rather broad. Tibiae straight, wide, flattened, slightly
longer than femora, their inner margin with small preapical enlargement,
without denticles. Tarsi three-segmented, considerably shorter than tibiae,
their first segment longer than broad and distinctly longer than second; third
segment equal in length to the preceding two together, slender. Claws simple,
slender, short.

Species composition. Monotypic genus.

Comparison. Mostclosely resembles the present-day genus Corticarina
Reitt., from which it differs in the transverse segments of antennae, absence
of a distinct pubescence on elytra, and five visible abdominal sternites.
Differs from the genus Melanophthalma Motsch. in the absence of femoral
lines on the first [abdomen] sternite and narrower scutellum without folds
and grooves; from Corticaria Marsh. it differs in the slightly serrate lateral
margin of pronotum and narrower scutellum; from Migneauxia Duval in the
11-segmented antennae, elongate body and longer first tarsal segment; from
Cortilena Motsch. and Diarthrocera Broun* in the three-segmented antennal
club.

Remarks. Succinimonitia definitely belongs to Lathridiidae. It is a very
typical representative of the family, fairly similar to the present-day forms,
and the combined characters relegate it to the rather advanced subfamily
Corticariinae. Ecologically Succinimontia hardly differs from the extant
Corticaria, Corticarina or Melanophthalma to which it is most closely
related. Representatives of this group inhabit shelters, particularly in the
forest litter and other heaps of plant detritus, sometimes under the bark of
dead wood, in burrows and nests of different animals, etc.; they feed on mold
fungi.

* Author’s name could not be confirmed—General Editor.

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This genus is the first Mesozoic representative of the family
Lathridiidae.

Succinimontia infleta Zherichin, sp. nov.
(Plate XII, Photo 4; Figure 75)

Species name coined from “infleta’ (Latin)}—unwept.

Holotype. No. 3130/31 , PIN, inclusion of almost entire beetle in
retinite, sex not determined. Taimyr national district, Khatanga region,
Yantardakh area on right bank of Maimechi river, 5 km upstream from its
mouth (Yantardakh site). Upper Cretaceous, ? Cognac-Santonian, Kheta
series.

Material. Holotype.

Description. Color brown, antennae and legs slightly lighter.

Interorbital width of frons twice the diameter of eyes, finely shagreened.
First antennal segment strongly clavate, its length twice its width, width of
second half the apical width of first and half its length, almost cylindrical,
very slightly broadening apically; ninth twice as long as eighth, ovo-trian-
gular, equal in length and width; tenth of same length and width as ninth,
rounded; eleventh 1.7 times longer than wide, parallel-sided, almost rec-
tilinearly tapering anteriorly in apical third.

Pronotum one-third wider than long, rounded laterally, without sharply
projecting corners, only with small denticles arranged as follows: two com-
paratively strong, pointed, triangular denticles in the anterior third of
pronotum, in its wide portion; one very small, smooth, rounded denticle
behind them; one comparatively strong, triangular denticle siightly behind
the middle. Beyond this rear denticle, the lateral margin of the pronotum is
only very indistinctly wavy. Width of anterior margin of pronotum three-
fourths its basal width. Pronotum with fine, indistinct shagreening.

Elytral shoulders almost one-third wider than pronotal base; sides of
pronotal disk smoothly rounded; lateral margin not flat. Elytra more convex
than pronotum. Rows of punctures rather distinct, regular; intervals between
them broad, flat, finely shagreened. Punctures of rows small, rounded.

Ventrally body uniformly, finely (slightly coarser than dorsally)
shagreened.

Dimensions. Body length 1.25 mm.

RHYNCHOPHORA

Beetles belonging to the Rhynchophora were for a long time considered an
independent taxon of a high rank; recently, however, they have been assigned

*Given as No. 3311/31 both in Fig. 75 and Plate XI, Photo 4—General Editor.

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Cc

: Q5mm

a
Fig. 75. Succinimontia infleta, sp. nov.; holotype PIN No. 3311/31: a—dorsal
view; b—antenna; c—margin of pronotum. Yantardakh, Upper Cretaceous.

to a superfamily of the infraorder Cucujiformia in which they constitute the
most advanced and youngest group. A study of the Mesozoic rhynchophoran
material gave a totally unexpected result. In the Late Jurassic, Rhynchophora
turned out to be the most diverse and abundant group among all Polyphaga.
Moreover, the recent finds of Triassic beetles hardly differing from the
Jurassic Eobelidae, compel us to consider them one of the most ancient
groups of the Polyphaga. For this reason here we have adopted a neutral
position and retained the name Rhynchophora without assigning them a
definite rank. Extant Rhynchophora are considered by the author to include
12 families: Belidae, Nemonychidae, Oxycorynidae, Aglycyderidae, Proter-
rhinidae, Antribidae, Urodontidae, Brenthidae, Attelabidae, Curculionidae,
Scolytidae, and Platypodidae.

The exceptionally valuable material on the Late Jurassic insects, col-
lected in the Karatau site of South Kazakhstan, has provided the basis for

evaluating the taxonomic position and organizational level of the Late

Jurassic Rhynchophora. Like many of the other groups of Coleoptera known
from Karatau, the Rhynchophora are already a highly specialized group for
which the initial evolutionary stages have long past. They are rich in species
and strongly differentiated. However, despite the high level of differentia-
tion, not one of the better preserved specimens examined could be assigned
to any of the modern rhynchophoran families, although a few of them exhibit
a tendency toward development along some of the same [evolutionary] paths.
In these instances it is possible to observe the intial stages of differentiation
towards the extant families.

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As a preliminary approach to processing the fossil material, the inves-
tigator compares it with modern representatives of rhynchophoran families
in order to ascertain whether the Mesozoic remains belongs with one of these,
or whether it is necessary to erect an independent taxon of familial rank.
Naturally, such comparisons should be made with families retaining the most
archaic features, such as the Belidae, Nemonychidae and Oxycorynidae
(Fig. 76).

The present-day Belidae—inhabiting Australia, South America, and
southern North America—possess many indisputably archaic features on top

Fig. 76. Head of primitive Rhynchophora; a, b—Diodirrhynchus austriacus
Oliv., family Nemonychidae, extant; c, d—Rhinotia sp., family Belidae, extant;
e, {—Oxycorynus melanocerus Sc., family Oxycorynidae, extant; g—Eobelus
sp., family Eobelidae, specimen PIN No. 2384/516; Karatau, Upper Jurassic.

143

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194

of which they have developed a few specialized characteristics. The
Australian genus Rhinotia should be considered the most primitive repre-
sentative among the recent species of this family, since its general appearance
is very similar to the Late Jurassic “‘belidoid” Rhynchophora. However, the
whole range of distinguishing features characteristic of Recent Belidae,
which are of basic diagnostic importance, differ from those established for
Jurassic forms. A low placement of the rostrum is characteristic of all belids;
the ventral surface of the rostrum is in line with the gular surface of the head
capsule, and ihe frontal surface of the head capsule forms an angle of up to
90° with the dorsal surface of the rostrum. This type of head structure is
peculiar to belids. The pronotum of extant belids has rounded margins,
without traces of the notopleural suture or of ridges. The pleurosternal sutures
are usually well developed. The antennae of belids lack a club, their first
segment is not enlarged, and they are attached at about midlength of the
rostrum in males, or at its base in females. The metathorax* is convex,
divided by a longitudinal groove. The abdominal sternites are more or less
equal in length. The legs are poorly specialized. The mandibles are of the
scraping type, with two to three denticles on the broad mesal edge.

A morphological comparison between most Jurassic “belidoid”
Rhynchophora and Recent forms shows the usual high degree of similarity
in addition to several fundamental differences. The head and rostrum are
similar in both groups: the rostrum always originates from the ventral margin
of the head capsule in the Jurassic as well as the Recent forms, but the apex
of the Jurassic rostrum is broad and noticeably (often strongly) compressed
dorsoventrally, whereas in the Recent forms it is cylindrical. The structure
of the Jurassic rostrum is very similar to that of primitive Nemonychidae
(Diodirrhynchus) or Attelabidae (Rhynchitinae). In Jurassic forms the man-
dibles were apparently always falcate with a single tip, and dorsoventrally
flattened as in Nemonychidae. The antennae of all the Jurassic species
examined resemble those of Recent Nemonychidae and Rhynchitinae; they
are attached in the middle or apical third of the rostrum but never basally.
Their first segment is not, or only slightly enlarged in comparison with the
following segments; the last three form a loose club. The pronota of all the
Late Jurassic “belidoids” have distinct notopleural ridges and probably a
corresponding suture or its traces which are absent in Recent belids. The legs
of the Jurassic forms are often more specialized than in the extant forms. The
abdominal sternites are always of nearly the same length. The comparison
thus shows that the Jurassic rhynchophorid beetles cannot be included in the
family Belidae.

*Since the metanolum is covered by the elytra, we may surmise that it is the ventral surface
of the metathorax which is indicated here, and presumably the metastemum itself. A ventral
perspective can certainly be assumed for the subsequent descriptions of the meso- and
metathoraces of fossil rhynchophorans preserved with closed elytra—Scientific Editor.

Ws)

The Jurassic forms far less strongly resemble the present-day Oxy-
corynidae. They are similar mainly in having a distinct notopleural ridge. In
the present-day oxycorynids the rostrum originates from the middle part of
the head capsule and its upper and lower sides form an obtuse angle with the
frontal and gular surfaces. In the Oxycorynidae, the antennae are always
attached at the base of the rostrum and generally have a rather distinct and
compact club which is often four-segmented. None of these characters is
known in the Jurassic forms, and consequently the latter cannot be included
in the family Oxycorynidae.

There are even fewer common structural features among the Jurassic
forms and the extant members of the families Nemonychidae and At-
telabidae. These mainly include structural similarities in the apex of the
rostrum, in the antennae and to some extent the abdomen. The similarities
are least between the Jurassic forms and the extant representatives of the true
snout-beetles, family Curculionidae.

Thus, a comparative analysis of the main external morphological fea-
tures of the fossil Rhynchophora shows that they do not belong to any of the
Recent families and represent an independent taxon of the familial rank.

The similarity of the main characters in all the Jurassic rhynchophorid
beetles studied by the author (with the exception of the controversial ones)
strongly suggests that they are the representatives of a single family and do
not belong to any of the Recent families. This conclusion is well supported
by the fact that, in a single specimen, characteristics restricted to the Jurassic
fauna occur in combination with a series of features which are individually
diagnostic of various representatives of the most primitive families which
are presently relictual. Hence the Jurassic Rhynchophora described here are
considered to be representatives of a now extinct special family Eobelidae,
whose several offshoots form independent families.

Family EOBELIDAE L. Arnoldi, Fam. Nov.

Diagnosis. Head rounded or transverse, convex frontally, steeply descending
to base of rostrum; rostrum disposed in lower part of head capsule so that
lower side of rostrum is in line with gular surface. Antennae without distinct
scape; first antennal segment not essentially different from other segements
or not more than twice as long as second segment. Antennae usually eleven-
segmented, with a loose, often indistinct club, attached at about midlength
of rostrum or, rarely, closer to its apex. Eyes always well developed,
noticeably shifted anteriorly. Pronotum always with distinct notopleural
ridge, abdomen with five similar sternites, evidently somewhat movable.
Structure of legs normal for superfamily, sometimes highly specialized.

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Description. Small- or medium-sized, elongate, slightly convex beetles,
apparently. with weakly sclerotized elytra. Head including rostrum, pro-,
meso- and metathorax as well as extremities highly sclerotized.

Head capsule laterally rounded, generally transverse, occipital foramen
wide. Eyes always well developed, not placed laterally but noticeably shifted
forward toward the frontal part of capsule, small or moderately large,
rounded. Frontal surface convex, generally projecting anteriorly and dorsal-
ly. Rostrum for the most part rather slender, always well developed, located
on lower part of head capsule, its lower side in line with gular surface, upper
side forming right angle or obtuse angle (90—130°) with frontal surface.
Length of rostrum highly variable, never less than the length of pronotum. It
[= rostrum] is straight or curved to a varying degree, curvature usually very
gentle. In most cases, rostrum flattened toward apex and broadened to a
varying degree, as in Nemonychidae (Diodirrhynchus) or Rhynchitinae. The
structure of mouth parts could not be studied in the usual detail, but in some
impressions their structure was rather close to the extant Nemonychidae
(Diodirrhynchus), i.e., mandibles with a singe apex and flattened dorso-
ventrally, without large denticles. Maxillae apparently free, not covered by
mentum and with highly reduced palps. Distinct gular sutures visible in good
impressions. Antennae generally attached near midlength of rostrum, rarely
close to its apex, and never at its base. A short, shallow pit is sometimes
noticeable at the place of antennal attachment and may represent a rudiment
of the future antennal socket. Antennae attached at the base of rostrum ina
single specimen, thick and strongly broadening toward apex, displaying
features of considerable specialization. In the same specimen, mandibles also
considerably enlarged, resembling those in males of some Recent Brentidae.
In all specimens in which antennae are well preserved, they are eleven-seg-
mented, with a loose, poorly differentiated three-segmented, club; none with
a marked elongation of the first segment, at most 1.5 to 2 times longer than
the following segments; second or third segment sometimes elongate, 1.e.,
the Jurassic forms still lacked the present-day scape and their antennae were
not geniculate.

Viewed laterally, prothorax always shows distinct division into notal
and sternal parts; notopleural ridge distinctly visible. A notopleural suture,
sometimes noticeable below the ridge, apparently also present. The sculptur-
ing On pronotum usually coarser than on pleura, with dense and rather coarse
punctation. Shape of pronotum rather uniform, generally transverse, weakly
rounded laterally, anteriorly weakly emarginate, basally straight or weakly
emarginate. Forecoxae always set part, generally by less than half their
diameter.

Elytra usually elongate, more or less parallel-sided, rarely uniformly and
weakly rounded. Elytra rarely short and broad. Punctate grooves, rows of
punctures or pits generally distinct. No hairs, scales or bristles visible,

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although they would have been visible [if present] on several of the high
quality impressions. Apices of elytra jointly or rarely separately rounded. In
one specimen each apex extended into a short triangular cusp. Lateral
(epipleural) margin of elytra usually slightly curved above hind coxae or
almost straight.

In most cases the mesothorax could not be examined in detail, neverthe-
less all its sutures were definitely present and the sclerites were differentiated.
Scutellum very rarely visible; middle coxae generally separated by a distance
nearly equal to their diameter, sometimes wider. Metasternum well
developed in all but one example, long usually 2.5—3 times longer than
diameter or coxae; episterna rectangular, sometimes with longitudinal suture
or groove.

Abdomen always with five similar sternites, more or less movable. Signs
of fusion between the first and second sternites or of a pronounced enlarge-
ment of the first sternite were not observed.

Structure of legs differs little from that in Recent families. In more
primitive forms, they have slightly dialated straight femora, slender straight
tibiae, and long narrow tarsi. In a number of cases, first [tarsal] segment
largest and broadest, third always bilobed, unguiculate segment narrow and
moderately long. In some Jurassic forms, the legs display highly specialized
features: femora strongly thickened, particularly the forefemora which are
sometimes thicker than long; tibiae slender, straight and short; tarsi excep-
tionally enlarged, almost equal in length to tibiae, but more than three times
the width of tibial apex, apparently covered by dense and soft setae since
their margins are highly diffuse in the impressions. There is evidence to
indicate that such tarsi are a secondary sexual character of males. In others
with normal femora, strongly curved tibiae or tibiae armed with apical spurs
are present.

Composition. Four subfamilies in the Late Jurassic of South Kazakh-
stan, described below.

Comparison. Resembles Belidae in the position of rostrum, the lower
side of which is continuous with the ventral surface of head, but differs from
it in the ridge and traces of a suture on the sides of pronotum, dorsoventral
flattening of the apex of rostrum and the frequently clavate antennae.
Resembles Oxycorynidae in the structure of prothorax; resembles Nemony-
chidae and Attelabidae (Rhynchitinae) in the structure of antennae and apex
of rostrum, but differs from them in the character of rostral base.

Remarks. The study of often incomplete and inadequately distinct im-
pressions of insects poses considerable difficulties in the taxonomic treat-
ment of the Mesozoic material. The impossibility of utilizing the whole range
of characters encountered in the extant forms, frequently impels one to study
only the most noticeable external features, making the conclusions less
reliable and occasionally tentative. The nature of the fossilization and sub-

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sequent deformation of the impressions under heavy pressure, hinders iden-
tification of a specimen’s species or genus; it is [therefore] possible that
specimens of a single species will be described under different genera.
Despite these difficulties, an experienced taxonomist usually finds sig-
nificant distinguishing characters to build a phylogenetically based system
of classification.

Subfamily Eobelinae L. Arnol’di Subfam. Nov.

Diagnosis. Rostrum generally slender and long, usually longer than head
capsule and pronotum together. Antennae attached at middle of rostrum or
near it. Pronotum small, dorsally flat, lateral ridges not sharp. Elytra usually
more or less parallel-sided or with a weakly rounded lateral margin, dorsally
flat or weakly convex. Abdomen with fairly similar sternites. Legs often
highly specialized, with distinctly thickened femora and enlarged tarsi. Hind
tibiae sometimes with large apical spur.

Composition. Three tribes in the Late Jurassic of South Kazakhstan.

Comparison. Most closely resembles the subfamily Oxycorynoidinae,
differs in the absence of sharp ridges on the sides of pronotum and usually
longer and thinner rostrum.

Tribe EOBELINI L. Arnol’di, trib. nov.

Diagnosis. Rostrum slender and long, considerably longer than head capsule
and pronotum together, almost straight or very slightly curved. Elytra flat or
weakly convex, more or less parallel-sided, epipleural margin slightly curv-
ing above hind coxae. Legs with slender straight femora and tibiae, or
specialized with highly thickened forefemora and very large foretarsi, or with
hind tibiae armed with large spurs.

Composition. Five genera in the Late Jurassic of South Kazakhstan
(Karatau).

Comparison. Distinguished from other tribes by the longer rostrum.

Genus Eobelus L. Arnol’di, gen. nov.

Type species. E. longipes, sp. nov. Upper Jurassic of South Kazakhstan.

Diagnosis. Most strongly resembles present-day Belidae in general
appearance. Head strongly convex in frontal part, with large eyes. Rostrum
long, almost cylindrical, slightly curved. Pronotum flat, with notopleural
ridge. Elytra more or less parallel-sided, flat. Legs long, with slender femora
slightly thickened toward middle, tibiae slender and straight, tarsi long and
narrow. Metathorax moderately convex.

Species composition. Monotypic genus.

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Comparison. Differs from other species in almost not thickened
forefemora and flat elytra.

Remarks. Based on general appearance, structure of legs, elytra and
head, this genus is apparently the most primitive of all the Jurassic
Rhynchophora studied by the author.

Eobelus longipes L. Arnoldi, sp. nov.
(Plate XII, Photo 5; Figure 77)

Species name coined from ‘longipes’ (Latin)—long-legged.

Holotype. No. 2452/275, PIN, almost entire impression of a beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley near Uspenskoe (Karatau-Galkino site). Upper
Jurassic, Karabastau series.

Material. Besides holotype, impression of a beetle lying on its back,
specimen No. 2384/516 from Karatau-Mikhailovka site (Fig. 76g) possibly
belonging to this species.

Description. Head rather larger, convex, frontal part sloping to base of
rostrum at an angle of about 80°. Eyes large, round, their lower margin not
reaching lower side of rostrum by about one-third the width of rostral base,
their length almost one-and-a-half times that of latter. Rostrum very slightly
curved, noticeably thin in apical third. Antennae noticeably shorter than
rostrum, with comparatively distinct club. Pronotum dorsally flat, distinctly
and rather prominently punctate on disk. Prothorax with pleurosternal suture.
Elytra weakly convex, apices acutely angled, 3.3 times longer than pronotum,
rather fine punctures visible on anterior half of disk. Metathorax long, but
comparatively weakly convex. Forelegs with almost nonclavate long femora,
longer than pronotum; tibiae even longer, narrow and almost straight, 1.75
times longer than rostrum. Foretarsi narrow. Middle femora shorter and
thinner than forefemora, tibiae straight and siender, hind femora not thinner
than middle and of same length, tibiae straight, tarsi slender and long. Third
bilobed tarsal segment small, shorter than remaining segments.

Dimensions. Body length 8.2 mm (here and throughout the remainder
of the text the body length does not include the rostrum); length of rostrum
3.6 mm.

Genus Archaeorrhynchus Martynov, 1926

Type species. A. tenuicornis Martynov, 1926; Upper Jurassic of South
Kazakhstan.

Diagnosis. Rostrum almost straight, noticeably thinning toward apex,
dorsoventrally flat at apex. Antennae with slightly elongate first three seg-
ments. Sides of pronotum with notopleural ridge. Forefemora strongly thick-

200

Fig. 77. Eobelus longipes, sp. nov.; holotype PIN No. 2452/275. Karatau,
Upper Jurassic.

ened, sometimes thicker than height of head capsule, foretarsi generally very
large (in males ?) and wide, particularly their first segment. Metathorax
convex.

Species composition. Four species in the Late Jurassic of South
Kazakhstan.

Comparison. Differs from all genera except Probelus in the strongly
thickened femora, differs from the latter in the straight foretibiae.

Remarks. The genus Archaeorrhynchus was established by Martynov
(1926) from the same Karatau site. The detailed description and good
illustration leave no doubt that Martynov described a representative of the
same genus as that which was present in the material examined by me.
However, Martynov’s description indicates the presence of a fine and long
shaft (scape) and an antennal groove in his A. tenuicornis Mart. This feature
was not observed in any of the well preserved antennae in the material at my
disposal. Even markedly elongate first segments have never been observed,
and it is definite that geniculate antennae did not yet exist in the Jurassic
forms, rather Martynov mistook the distinct impression of the flagellum for
the scape, and the two lateral margins of the basal part of the rostrum for the
antennal grooves.

Archaeorrhynchus acutirostris L. Arnoldi, sp. nov.
(Figure 78)

Species name coined from ‘acutis’ (Latin)—sharp, and ‘rostrum’ (Latin)—
beak.

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201

Holotype. No. 2554/715, PIN, almost entire impression of a beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head rather large, with frontal surface steeply descending
from temple to rostrum, forming almost right angle. Eyes comparatively
small, round. Rostrum rather slender, basally thinner than length of eyes,
markedly thinning toward apex, almost subulate (in lateral position !), in
apical half slightly curving from point of antennal attachment. Antennae
attached almost at middle of rostrum, their structural details indistinct in basal
half, but undoubtedly lacking a long scape; club weakly differentiated. Lower
side of rostrum slightly convex from base, asa resulta slight gular indentation
is observed. Pronotum transverse with basal arcuate emargination. Elytra
apparently elongate-oval, their apical angles acute, faint grooves discernible
on disk, lateral margins almost straight. Metathorax rather convex, [ventral-
ly] almost twice as long as width of hind coxae. Abdomen with subequal
sternites, third being the shortest. Specimen apparently female since ab-
domen extends posteriorly into a short tube. Forelegs considerably larger
than others, their femora strongly thickened, tibiae slender, tarsi not
preserved. Middle femora approximately half the thickness of forefemora,
tibiae straight, tarsi large and long. Hind legs shorter than others, hind femora
somewhat thicker than middle femora and two-thirds their length; tibiae
straight, tarsi comparatively small.

Dimensions. Body length 7.8 mm, length of rostrum 3.2 mm.

Comparison. Distinguished by hind femora which are much shorter than
middle femora.

Fig. 78. Archaeorrhynchus acutirostris, sp. nov.; holotype PIN No. 2554/715.
Karatau, Upper Jurassic.

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Archaeorrhynchus paradoxopus L. Arnol’di, sp. nov.
(Plate XII, Photo 6; Figure 79)

Species name coined from ‘paradoxa’ (Greek)—strange, and ‘pous’
(Greek)—foot.

Holotype. No. 2335/42, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Uspenskoe settlement (Karatau-Galkino site). Upper
Jurassic, Karabastau series.

Material. Holotype.

Description. Head moderately large, short; frontal surface forming an
almost right angle with rostral base. Eyes moderately large, longitudinally
slightly oval, shifted anteriorly. Rostrum long, basally narrower than length
of eyes, strongly thinning distally, subulate, almost straight. Antennae at-
tached in basal half of rostrum, their first segment longer than remaining
segments, club present but poorly differentiated. Total antennal length almost
equal to length of rostrum. Pronotum dorsally flat; length of rostrum 2.3 times
greater than pronotal length along upper surface. Punctation fine, dense.
Elytra slightly convex, posteriorly abruptly rounded off, rows of large
punctures distinctly visible on disk. Length of elytra three times that of
pronotum. Epipleural margin straight. Prothorax small, forecoxae shifted
closer to its posterior margin. Metathorax long, more than three times longer
than width of hind coxae, moderately convex, metepisterna broad, fine
transversely rugose punctation distinctly visible. Abdomen with similar
sternites, apically rounded. Forelegs with very strongly thickened femora,
femoral thickness three times that of rostral base. Foretibiae comparatively
slender and straight. Tarsi very large, almost as long as tibiae and more than
two-and-a-half times wider than them. Their structural details could not be
examined; apparently they are densely covered by fine pubescence, and are
structurally similar to the tarsi of A. acutirostris, sp. nov. Middle femora
weakly clavate, with fine transverse striations; tibiae long and straight; tarsi
slender. Hind femora thinner and shorter; tibiae straight, short, apically
apparently with tuft of bristles; tarsi slender.

Dimensions. Body length 10.2 mm, length of rostrum 5.4 mm.

Comparison. Resembles A. /atitarsus in the structure of foretarsi, differs
from it in smaller head.

Archaeorrhynchus latitarsus L. Arnol’di, sp. nov.
(Figure 80)

Species name coined from ‘latus’ (Latin)—wide, and ‘tarsus’ (Latin)—foot.
Holotype. No.2239/531, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-

eae ae ol ae

149

203

Fig. 79. Archaeorrhynchus paradoxopus, sp. nov.; holotype PIN No. 2335/42.
Karatau, Upper Jurassic.

karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head massive, frontal surface rather steeply descending to
base of rostrum, forming an angle of about 50° with its upper surface. Eyes
round, small, their lower margin level with upper side of rostrum. The latter
relatively thick, more than length of eyes, uniformly narrowing toward apex
and very slightly curved. Antennae attached at end of basal third of rostrum,
slender, with non-abrupt club. First antennal segment longest; second ap-
proximately one-third and third only slightly shorter than first, following

Fig. 80. Archaeorrhynchus latitarsus, sp. nov.; holotype PIN No. 2239/531.
Karatau, Upper Jurassic.

151

204

segments less than half of third. Total antennal length almost equal to that of
head including rostrum, segments of club triangular, last segment rhom-
boidal. Pronotum short, posteriorly rounded, with distinct notopleural ridge,
dorsally weakly convex. Elytra apparently more or less parallel-sided, angu-
larly tapering apically. Epipleural margin of elytra straight. Prothorax rather
high [= tall], forecoxae closer to its posterior margin. Mesothorax short, with
distinct sutures between sclerites, middle coxae large. Metathorax very long,
3 times longer than width of hind coxae, uniformly convex, epimera rectan-
gular. Abdomen with similar segments, longer than elytra. Forelegs with
strongly thickened femora, their width 3 times the basal thickness of rostrum.
Foretibiae rather slender, almost straight; tarsi strongly dialated, their first
segment two-and-a-half times broader than tibial apex, apically emarginate;
second segment anteriorly tapering, one-third the length of first; third very
small; unguiculate segment short and apparently slender. Middle femora
slightly thickened. Hind legs with gently clavate femora; slender, straight
tibiae, and slender tarsi.

Dimensions. Body length 8.8 mm, length of rostrum 3.8 mm.

Comparison. Differs from closely related A. paradoxopus in the more
massive head, and sloping frons.

Genus Probelus L. Arnol’di, gen. nov.

Genus name coined from genus Belus.

Type species. P. curvispinus, sp. nov. Upper Jurassic of South Kaza-
khstan.

Diagnosis. Head large or moderately large; rostrum almost straight,
longer than head capsule and pronotum together. Antennae slender, with
distinct club, slightly shorter than rostrum, attached before its middle.
Pronotum short, transverse, with sharp notopleural ridge, dorsally flat. Elytra
more or less parallel-sided, dorsally flat. Foremora slightly or moderately
clavate, tibiae slightly curved, tarsi small or long, but narrow. Hind tibiae
having mild preapical depression on inner side, with large straight or arched
spur on anterior apical angle.

Species composition. Three species in the Late Jurassic of South
Kazakhstan.

Comparison. Differs from other genera in hind tibiae with depression
on inner side and large apical spur.

Probelus curvispinus L. Arnoldi, sp. nov.
(Plate XIII, Photo 1; Figure 81)

Species name coined from ‘curvus’ (Latin)—curved, and ‘spina’ (Latin)—
point, spur.

152

205

Fig. 81. Probelus curvispinus, sp. nov.; holotype PIN No. 2554/709. Karatau,
Upper Jurassic.

Holotype. No. 2554/709, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Holotype. .

Description. Head moderately large, frontally convex, forming right
angle with base of rostrum. Eyes small, somewhat larger than basal width of
rostrum, rounded; rostrum almost straight, strongly tapering toward apex;
antennae attached at beginning of anterior third of rostrum, their first segment
not enlarged. Pronotum with weakly raised posterior margin. Elytra almost
flat, 3.3 times longer than pronotum, epipleural margin scarcely curved. Legs
short, all femora rather strongly clavate, particularly forefemora. All tibiae
curved; fore and middle tibiae short; hind tibiae longer, depressed preapical-
ly, apical angle extending into falcate spur, directed anteriorly and ventrally.
Foretarsi small, structure typical, hind tarsi narrow but apparently long.

Dimensions. Body length 7.0 mm, length of rostrum 3.2 mm.

Comparison. Differs from other species in more strongly thickened
femora.

Probelus longitarsus L. Arnol’di, sp. nov.
(Figure 82)

Species name coined from ‘longus’ (Latin)—long, and ‘tarsus’ (Latin)—
foot.

Holotype. No. 2066/2811, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

153

206

Fig. 82. Probelus longitarsus, sp. nov.; holotype PIN No. 2066/2811. Karatau,
Upper Jurassic.

Material. Holotype.

Description. Head small, frontal surface forming angle of about 100°
with upper side of rostrum. Eyes small, their lower margin almost level with
upper side of rostrum. The latter slender, moderately tapering toward apex,
somewhat more noticeably curved preapically, its length 1.3 times that of
head capsule and pronotum together. Antennae attached immediately before
rostral middle, slender and rather long, their details indistinct. Pronotum very
slightly convex dorsally, its posterior margin slightly upcurved. Elytra weak-
ly convex, acute angled apically, epipleural margin slightly emarginate
above hind coxae. Metathorax long, [ventrally] convex. Legs with moderate-
ly thickened femora, middle tibiae noticeably curved in basal fourth, and
smoothly depressed preapically on inner side. Hind tibiae weakly curved
anteriorly, with broad and gently sloping preapical depression on inner side,
apical spur short, curved. Foretarsi poorly preserved, but undoubtedly nar-
row; middle and hind tarsi narrow and long, only slightly shorter than tibiae.

Dimensions. Body length 8.0 mm, length of rostrum 3.2 mm.

Comparison. Differs from type species in possessing less thickened
femora, from P. tibialis in curved spurs of hind tibiae.

Probelus tibialis L. Arnoldi, sp. nov.
(Figure 83)

Species name coined from ‘tibia’ (Latin)—shin.

Holotype. No. 965/45, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Uspenskoe settlement (Karatau-Galkino site). Upper
Jurassic, Karabastau series.

Material. Holotype.

207

Description. Head large, transverse; eyes large, strongly shifted for-
ward. Rostrum rather wide, with distinct though slight constriction at junc-
ture with antennae, i.e., apex of rostrum slightly broad at commencement of
apical third. Antennae rather short, third segment longest, club distinct.
Pronotum anteriorly weakly emarginate, almost straight basally, sides
moderately rounded. Elytra longitudinally oval, parallel-sided in anterior
half. Forecoxae strongly convergent, not contiguous, middle coxae also
slightly separated, hind coxae separated by half their length. Meso- and
metathorax [ventrally] with longitudinal groove. Length of metathorax twice
the width of hind coxae. Legs with slightly clavate femora and rather thick,
very slightly curved tibiae; hind tibiae notched, apex of spur straight; tarsi
slender and short.

Dimensions. Body length 7.3 mm, width 2.6 mm; length of rostrum 1.9
mm.

Comparison. Differs from type species in possessing slightly [as ap-
posed to greatly] thickened femora, and from P. longitarsus in straight spurs
of hind tibiae.

ST

Fig. 83. Probelus tibialis, sp. nov.; holotype PIN No. 965/45; Karatau, Upper
Jurassic.

154

208

Remarks. This species is included in the genus Probelus on the basis of
the structure of the hind tibiae and general proportions of the body. Com-
parison with other species is not possible since the fossil is positioned on its
back.

Genus Probelopsis L. Arnoldi, gen. nov.

Genus name coined from genus Belus.

Type species. P. acutiapex, sp. nov. Upper Jurassic of South Kazakh-
stan.

Diagnosis. Head small; rostrum almost straight, slightly curved only in
apical third. Eyes large, their width 1.4 times the basal width of rostrum.
Elytra flat, apparently parallel-sided, their apices weakly drawn out into short
triangular processes. Legs with moderately clavate femora, forefemora
slightly thicker than remaining femora, tibiae rather slender, fore- and middle
tibiae slightly curved, hind tibiae straight. Inner angles of fore- and hind tibiae
with short spurs.

Species composition. Monotypic genus.

Comparison. Differs from the preceding two genera in the less thick-
ened femora, differs slightly from the first* in the drawn out elytral apices,
and from the remaining genera in the almost straight rostrum.

Probelopsis acutiapex L. Arnoldi, sp. nov.
(Plate XIII, Photo 2; Figure 84)

Species name coined from ‘acutus’ (Latin)—pointed, and ‘apex’ (Latin)—
tip.

Holotype. No. 2239/1554, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head not high**, eyes large, round, rostrum long. Anten-
nae attached at two-fifths the length of rostrum from base. Rostrum longer
than head capsule and pronotum together. Pronotum rather short, dorsally
noticeably convex, notopleural ridge distinct, forecoxae at posterior margin
of prothorax. Elytra slightly convex dorsally, anteriorly truncate, weakly
emarginate on epipleural margin. Elytral apices drawn out into short points.
Legs rather long, femora very weakly clavate, forefemora thicker, their

*In this context the term ‘first’ probably refers to the genus Eobelus which is the first genus
listed in the tribe Eobelini—Scientific Editor.

“As used here, the term ‘high’ does not refer to the orientation or position of the head, but
to the length of its dorsoventral axis—Scientific Editor.

209

Fig. 84. Probelopsis acutiapex, sp. nov.; holotype PIN No. 2239/1554; Karatau,
Upper Jurassic.

thickness almost twice the basal width of rostrum. Foretibiae linear up to
apical fourth and here noticeably curved, middle tibiae weakly and uniformly
curved, hind tibiae almost straight. Spurs on fore- and hind tibiae resemble
a short spine, directed anteriorly. Tarsi almost not preserved, but were
apparently slender and short.

Dimensions. Body length 7.3 mm, length of rostrum 2.8 mm.

Genus Belonotaris L. Arnol’di, gen. nov.

Genus name coined from genera Belus and Notaris.

Type species. B. punctatissimus, sp. nov.; Upper Jurassic of South
Kazakhstan.

Diagnosis. Head small, rostrum long, always uniformly and smoothly
curved; antennae attached at middle. Elytra noticeably convex, usually with
rows of punctures, with distinct emargination of epipleurai margin above
hind coxae. Legs with slightly thickened femora, tibiae lacking armature, and
small tarsi.

Species composition. Three species (one included tentatively) in the
Jurassic of South Kazakhstan.

Comparison. Distinguished by convex elytra with distinct emargination
of epipleural margin, uniformly curved rostrum and tibiae lacking armature.

155

210

Belonotaris punctatissimus L. Arnoldi, sp. nov.
(Plate XIII, Photo 3; Figure 85)

Species name coined from ‘punctum’ (Latin)—point.

Holotype. No. 2452/422, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Karabastau area (Karatau-Karabastau site). Upper Juras-
sic, Karabastau series.

Material. Holotype.

Description. Head small, frontal surface smoothly descending to base
of rostrum, forming obtuse angle (about 120-130") with it. Eyes small, round.
Rostrum slender, long, smoothly curved, its length somewhat more than total
length of head capsule plus pronotum. First three segments of antennae small,
cylindrical. Pronotum rather long, dorsally weakly convex, with dense and
coarse punctation. Elytra elongate, moderately convex, with rather strongly
curved epipleural margin, with distinct rows of large punctures, probably
forming a honeycombed pattern. Metathorax weakly convex, abdomen with
similar sternites. All femora moderately clavate, 1.7 times thicker than base
of rostrum; tibiae poorly preserved, but apparently straight and comparative-
ly slender; tarsi not preserved.

Dimensions. Body length 6.2 mm, length of rostrum 2.2 mm.

Comparison. Distinguished by longer pronotum.

Fig. 85. Belonotaris punctatissimus, sp. nov.; holotype PIN No. 2462/422';
Karatau, Upper Jurassic.

*Given as 2452/422 in the Text and Plate XII[I—General Editor.

156

211

Fig. 86. Belonotaris lineatipunctatus, sp. nov.; holotype PIN No. 2066/3209;
Karatau, Upper Jurassic.

Belonotaris lineatipunctatus L. Arnol’di, sp. nov.
(Figure 86)

Species name coined from ‘linea’ (Latin)—line, and ‘punctum’ (Latin)—
point.

Holotype. No. 2066/3209, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head small, frontal surface sloping to form obtuse angle of
120-30° with upper side of rostrum; eyes round, their diameter 1.2 times the
basal width of rostrum. Rostrum slender, uniformly weakly curved, its length
somewhat less than total length of head capsule and pronotum. Antennae
attached at about middle of rostrum, somewhat closer to its apex. They were
poorly preserved but definitely without a distinctly elongate first segment.
Pronoium short, weakly convex, dorsally with fine and dense punctation;
elytra noticeably convex, with punctate grooves, punctures in them large,
forming a honeycomb pattern. Lateral margins of elytra strongly curved
(possibly as a result of distortion of the specimen during fossilization).
Abdomen with identical sternites, posteriorly pointed. Legs rather slender
and long, forefemora more strongly clavate than remaining femora. Tibiae
almost straight, tarsi small and slender.

Dimensions. Body length 2.9 mm, length of rostrum 1.2 mm.

Comparison. Distinguished by short pronotum.

Belonotaris karatavicus L. Amol’di, sp. nov.
(Figure 87)

Species named after Karatau.

157

156

212

Holotype. No. 2066/2552, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head small, frontal surface initially more gently sloping,
then sharply descending almost at right angles to base of rostrum. Eyes round,
rather large, slightly larger than basal width of rostrum. The latter long, 2.25
times longer than pronotum, moderately thick, weakly and uniformly curved,
gradually thinning toward apex. Antennae attached just anterior to middle,
slender, with distinct three-segmented club; antennal length somewhat less
than that of rostrum. Pronotum a little convex, twice as long as head capsule.
Elytra moderately and uniformly convex, apices of angles acute, epipleural
margin noticeably emarginate above hind coxae. Metathorax comparatively
short, less than twice as long as width of hind coxae. Abdomen considerably
projecting beyond elytral apex. Legs with weakly clavate femora; tibiae
slightly curved, slender; tarsi poorly distinguishable but undoubtedly small.

Dimensions. Body length 8.5 mm, length of rostrum 4.0 mm.

Comparison. Differs from other species in the shape of head whose
frontal surface forms almost right angle with rostrum.

Remarks. Inclusion of this species in the genus Belonotaris raises some
doubts since the body proportions do not fully coincide with those of the two
preceding species. However, the shape of rostrum and elytra, and also the
structure of legs are very similar to those in the aforementioned species.

Tribe PROCURCULIONINI L. Arnoldi, trib. nov.

Diagnosis. Rostrum long, thick, distinctly narrowing at antennal junction,
apical part somewhat broadened. Mandibles large. Length of pronotum

Fig. 87. Belonotaris karatavicus, sp. nov.; holotype PIN No. 2066/2552;
Karatau, Upper Jurassic.

NS)

exceeding its width by less than two times, apical margin projecting at
middle. Elytra parallel-sided. Forefemora thickened.
Composition. Single genus in the Late Jurassic of South Kazakhstan.
Comparison. Differs from Eobelini in short thick rostrum, and elytra
without emargination of lateral margin; from Eccoptothoracini in the longer
pronotum.

Genus Procurculio L. Arnol’di, gen. nov.

Genus name coined from genus Curculio.

Type species. P. fortipes, sp. nov. Upper Jurassic of South Kazakhstan.

Diagnosis. Rostrum narrowest at beginning of anterior third, its preapi-
cal part broad, parallel-sided. Mandibles almost as long as apical width of
rostrum. Pronotum strongly rounded laterally, its anterior margin very slight-
ly projecting at middle.

Species composition. Monotypic genus.

or

Qty

=
= o a
i)
e Nae .
Qi
= ——
SS Se

Fig. 88. Procurculio fortipes, sp. nov.; holotype PIN No. 2066/2339; Karatau,
Upper Jurassic.

158

214

Procurculio fortipes L. Arnol’di, sp. nov.
(Plate XIII, Photo 4; Figure 88)

Species name coined from ‘fortis’ (Latin)—strong, and ‘pes’ (Latin)—foot.

Holotype. No. 2066/2339, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-westem flank
of Kashkarata river valley, Aulic area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head capsule almost semicircular, frontally steeply des-
cending to base of rostrum, which is somewhat shorter than head capsule and
pronotum together, gradually narrowing up to anterior third where antennae
are attached, then broadening sharply for a short distance, but further on up
to apex again parallel-sided. Base of rostrum only slightly narrower than the
interorbital width of frons. Antennae slender, their second and third segments
longer than remaining segments, club not abrupt; antennal length equal to
rostrum and head capsule together. Pronotum transverse, laterally rounded,
maximum width behind middle, appreciably narrower at posterior margin
than at anterior. Elytra 2.5 times longer than pronotum and only 1.2 times
broader, parallel-sided up to apical third, apices together broadly rounded.
Metathorax long; abdomen somewhat protruding beyond elytral apices. Legs
long; forefemora shorter than others, strongly swollen; middle and hind
femora slightly thickened, almost straight; tarsi apparently narrow.

Dimensions. Body length 7.5 mm, length of rostrum 2.4 mm.

Tribe ECCOPTOTHORACINI L. Arnol’di, trib. nov.

Diagnosis. Rostrum rather short, moderately thick. Pronotum strongly
transverse, its length less than half its width at base, posterior angles acute.
Anterior margin smoothly emarginate, emarginate basally [too]. Abdomen
with first three sternites identical, fourth considerably narrower with strongly
rounded emargination at posterior margin, fifth blunt. Legs not thickened.

Composition. Single genus in the Late Jurassic of South Kazakhstan.

Comparison. Distinguished by strongly transverse pronotum, tapering
anteriorly.

Genus Eccoptothorax L. Arnol’di, gen. nov.

Genus name coined from ‘eccopte’ (Greek)—emarginate, and ‘thorax’
(Greek)—chest.

Types species. E. latipennis, sp. nov. Upper Jurassic of South Kaza-
khstan.

ANS)

Diagnosis. Rostrum tapering from base to place of antennal attachment
slightly anterior to its middle. Length of pronotum two-fifths its basal width,
anterior margin noticeably narrower than posterior, latter deeply emarg:nate.
Metathorax long.

Species composition. Monotypic genus.

Eccoptothorax latipennis L. Arol’di, sp. nov.
(Plate XIII, Photo 5; Figure 89)

Species name coined from ‘latus’ (Latin}—broad, and ‘penna’ (Latin)—
wing.

Holotype. No. 2554/720, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head short and wide, rostrum moderately long, longer than
head capsule and pronotum together, moderately thick, apical part broader

Fig. 89. Eccoptothorax iatipennis, sp. nov.; holotype PIN No. 2554/720;
Karatau, Upper Jurassic.

159

216

than base. Antennae rather long, not shorter than head with rostrum. Their
first four segments slightly larger than remaining segments, club distinct.
Mandibles rather long. Pronotum laterally broadened from anterior to
posterior margin, both margins noticeably emarginate. Abdomen atypical of
Jurassic Rhynchophora: its sternites gradually decreasing in size from first
to fourth, which is strongly rounded and emarginate at posterior margin,
while the fifth set into the notch of fourth by almost half its length, posteriorly
triangular. Elytra rather wide, almost parallel-sided, probably convex in
posterior half. Broad, turned-under elytral epipleura may be the result of
flattening. Femora weakly clavate, rather long; tibiae poorly visible, but
apparently not thick, slightly curved or straight. Tarsi not visible.

Dimensions. Body length 6.5 mm, length of rostrum 1.8 mm, elytral
width in anterior half 2.6 mm.

Subfamily Oxycorynoidinae L. Arnol’di, Subfam. Nov.

Diagnosis. In general appearance very closely resembles Oxycorynidae.
Rostrum moderately long, antennae attached at its middle. Pronotum with
sharp longitudinal lateral ridge. Abdomen with rather similar sternites. Legs
usually rather short, sometimes with tibiae distinctly curved or armed with
short spurs.

Composition. Three tribes in the Late Jurassic of South Kazakhstan.

Composition. Resembles subfamily Eobelinae, differs in distinct lon-
gitudinal ridge of prothorax.

Tribe OXYCORYNOIDINI L. Arnol’di, trib. nov.

Diagnosis. Small or minute compact beetles; in most cases [length of
rostrum ]* not exceeding the length of head capsule and pronotum together,
rarely somewhat longer. Rostrum usually nearly straight, more rarely
moderately curved, usually slightly broadening at apex. Antennae always
attached near rostral middle. Position of rostrum on head capsule typical of
belidoids, but sometimes (if not owing to distortion during fossilization) on
the lower side, not fully in line with the gular surface. Elytra weakly convex
or flat, rounded or pointed apically, with acute sutural angle. Abdomen with
rather similar sternites. Legs usually small, generally lacking specialized
features; tibiae sometimes curved or armed with small spurs.

Composition. Four genera in the Late Jurassic of South Kazakhstan, of
which two (Ampliceps and Paroxycorynoides) stand somewhat apart and
probably merit separation into independent taxa of tribal rank. However, the

*The bracketed part of this phrase was extrapolated from the context; it was apparently
accidentally omitted in the Russian original—Scientific Editor.

Dei

data are insufficient for such a separation, and hence they are included in this
tribe.

Comparison. Distinguished from other tribes by the normal structure of
legs and antennal attachment close to midrostrum.

Genus Oxycorynoides L. Arnol’di, gen. nov.

Genus name coined from genus Oxycorynus.
Type species. O. similis, sp. nov. Upper Jurassic of South Kazakhstan.
Diagnosis. Comparatively short, compact, minute beetles. Head
moderate sized, rarely large, high*; eyes round, generally small. Length of
rostrum somewhat less than combined length of head capsule and pronotum,

_ rarely slightly more; rostrum straight or slightly curved, slightly tapering

160

toward apex (appears to be broadening apically in lateral, dorsal or ventral
views), antennae generally attached somewhat anterior to its middle.
Pronotum short, weakly transverse, with posterior angles rounded, noto-
pleural ridge present. Elytra weakly convex, apically with acute sutural angle
or more rarely rounded. Legs usually short, lacking Spociaiecd features,
tibiae straight, tarsi always slender, small.

Species composition. Six species (one tentatively included in this genus)
in the Late Jurassic of South Kazakhstan.

Comparison. Distinguished by slightly thickened femora and straight
tibiae.

Oxycorynoides similis L. Arnoldi, sp. nov.
(Plate XIII, Photo 6; Figure 90)

Species name coined from ‘similis’ (Latin)—similar.

Holotype. No. 2554/713, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailo-
vka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head moderately large; eyes slightly less than half the
height of head capsule at pronotal margin, round, positioned against rostral
base. Frontal surface forming obtuse angle (about 130°) with base of rostrum,
gular surface not coplanar with lower side of rostrum, forming an angle of
about 140—150° with it. Length of rostrum somewhat less than combined
length of head capsule and pronotum. Rostrum almost straight, slightly
thinning toward apex, antennae attached at its midlength. Pronotum weakly
convex, short. Elytra weakly convex in posterior half, with acute apical angle.

*See our comment on page 208—Scientific Editor.

218

Fig. 90. Oxycorynoides similis, sp. nov.; holotype PIN No. 2554 / 713; Karatau,
Upper Jurassic.

Epipleural margin of elytra almost straight. Rows of punctures on sides of
elytra. Metathorax long, posteriorly more convex [in ventral view]. Ab-
domen with similar sternites, terminally pointed, projecting beyond elytral
apex. Legs slender, short, tibiae apparently straight, tarsi not preserved.
Dimensions. Body length 3.0 mm, length of rostrum 0.8 mm.
Comparison. Distinguished by antennal attachment close to mid-
rostrum.

Oxycorynoides brevipes L. Arol’di, sp. nov.
(Figure 91)

Genus name coined from ‘brevis’ (Latin)—short, and ‘pes’ (Latin)—foot.
Holotype. No. 2239/1498 , PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank

Fig. 91. Oxycorynoides brevipes, sp. nov.; holotype PIN No. 2239/1489;
Karatau, Upper Jurassic.

*Given as 2239/1489 in Figure 91—General Editor.

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of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head with noticeably convex frontal surface, forming an
angle of approximately 100° with rostrum. Eyes rather large, positioned
against base of rostrum, their width considerably more than basal thickness
of rostrum. Rostrum scarcely shorter than pronotum, rather thick, slightly
narrowing toward apex, weakly and uniformly curved, antennae attached
slightly anterior to its middle, their first five segments small, remaining
segments not visible. Pronotum anteriorly projecting, dorsally very slightly
convex, its base slightly raised medially. Elytra dorsally flat, rounded apical-
ly; epipleural margin almost straight, length two and a half times greater than
that of pronotum. Rows of punctures visible on the sides of elytra
(honeycomb pattern ?). Abdomen with similar sternites, last sternite bluntly
rounded. Metathorax long, [ventrally] moderately convex posteriorly.
Forelegs somewhat larger than remaining legs, femora slightly clavate,
particularly middle and hind femora, tibiae straight and slender, tarsi small.

Dimensions. Body length 3.2 mm, length of rostrum 1.2 mm.

Comparison. Distinguished by slightly anteriorly projecting pronotum.

Oxycorynoides rohdendorfi L. Arnol’di, sp. nov.
(Figure 92)

Species named after B.B. Rohdendorf.

Holotype. No. 2239/1501, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Fig. 92. Oxycorynoides rohdendorfi, sp. nov.; holotype PIN No. 2239/1501;
Karatau, Upper Jurassic.

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Description. Head small, with slightly convex frontal surface and rather
small, noticeably anteriorly shifted eyes. Diameter of eyes less than basal
width of rostrum. Rostrum rather thick, almost straight, its length 1.7 times
that of pronotum, slightly thinning toward tip. Antennae attached a little
anterior to midrostrum, slender; basal segments slightly varying in length;
remaining segments poorly preserved, however a non-abrupt club distin-
guishable. Antennal apices reaching middle of pronotum. Pronotum small,
anteriorly truncate, disk slightly depressed dorsally (result of fossilization ?),
punctation dense and coarse. Elytra slightly convex dorsally, pointed apical-
ly, without noticeable sculpture. Metathorax long, [ventrally] slightly con-
vex. Abdomen with similar sternites, fifth sternite covered by pygidium.
Epipleural margin of elytra not curving above hind coxae, but uniformly
rounded. Legs poorly preserved, but their structure undoubtedly normal, hind
tibiae straight. Femora weakly clavate.

Dimensions. Body length 2.1 mm, length of rostrum 1.0 mm.

Comparison. Distinguished by anteriorly truncate pronotum.

Oxycorynoides ponomarenkoi L. Arnoldi, sp. nov.
(Figure 93)

Species named after A.G. Ponomarenko.

Holotype. No. 2066/2742, PIN, almost entire impression of beetie.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head relatively large, round; eyes small, round, noticeably
anteriorly shifted; rostrum straight, almost of uniform thickness, very slightly
broadening only at tip, apparently cylindrical. Antennae attached slightly
anterior to rostral middle, their total length scarcely more than that of rostrum,

Fig. 93. Oxycorynoides ponomarenkoi, sp. nov.; holotype PIN No. 2066/2742;
Karatau, Upper Jurassic.

ee oe er

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club distinct, first segment not enlarged. Pronotum slightly emarginate.,
posteriorly somewhat angularly projecting, disk with dense and fine puncta-
tion, laterally very weakly rounded. Elytra elongate-oval, posteriorly
pointed, smoothly rounded laterally, without noticeable bend above hind
coxae. [Ventral surface of] thorax not visible. Abdomen indistinctly visible,
however having similar sternites, slightly projecting beyond elytral apex, and
pointed at apex. Legs slender, femora long, slightly clavate; tibiae slender,
straight; tarsi not visible.

Dimensions. Body length 3.9 mm, length of rostrum 1.0 mm.

Comparison. Distinguished by anteriorly emarginate and posteriorly
projecting pronotum.

Oxycorynoides zherichini L. Amol’di, sp. nov.
(Figure 94)

Species named after V.V. Zherikhin.

Holotype. No. 2239/1555, PIN, almost entire impression of beetle.
South Kazakhstan. Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Fig. 94. Oxycorynoides zherichini, sp. nov.; holotype PIN No. 2239/1555;
Karatau, Upper Jurassic.

*Probably referring to the anterior margin. See comparison section —Scientific Editor.

63

2

Description. Head round, with small eyes placed closer to upper and
anterior side of head capsule. Frontal surface apparently sharply descending
to base of rostrum since head capsule is anteriorly bounded by arch, forming
a small hood over the rostral base. Rostrum parallel-sided almost up to apex,
moderately slender and angularly broadening only at apex itself. Its thickness
equal to width of frons between the eyes. Antennae attached at midrostrum,
slender, with scarcely enlarged first segment, and non-abrupt club, their
length noticeably greater than rostrum. Pronotum transverse, slightly emar-
ginate anteriorly and posteriorly, slightly rounded laterally. Elytra rather
short and wide, apically right angled, laterally almost parallel-sided. [Ventral
surface of] thorax not visible, abdomen slightly projecting beyond apex of
elytra, pointed. Legs with comparatively slightly thickened, short femora. As
far as visible, tibiae slender and straight. Tarsi not visible.

Dimensions. Body length 3.0 mm, length of rostrum 1.0 mm.

Comparison. Distinguished by anteriorly and posteriorly emarginate
pronotum, and short, wide elytra.

Remarks. Holotype lying on venter, hindering its comparison with other
species.

Oxycorynoides progressivus L. Arnol’di, sp. nov.
(Figure 95)

Species name coined from ‘progressio’ (Latin)—progress.

Holotype. No. 2066/2317, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Fig. 95. Oxycorynoides progressivus, sp. nov.; holotype PIN No. 2066/2317;
Karatau, Upper Jurassic.

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Description. Head rather large; eyes round, placed closer to lower
margin of head capsule. Rostrum not in same plane with lower surface of
head capsule but forming a very obtuse angle with it. It is difficult to decide
whether it is really so, or due to some twisting in the position of the head of
this specimen that it [the rostrum 7 is visible both in dorsal and lateral views.
Rostrum relatively short, slender in front of base, thinner than diameter of
eyes, gradually broadening toward apex. Antennae attached in anterior third
of rostrum, their basal segment scarcely larger than following segments;
segments gradually increasing in thickness and transformation to club not
abrupt. Pronotum with distinct anterior angles and weakly rounded sides,
base almost straight. Elytra comparatively short, only 2.2 times longer than
pronotum, their epipleural margin slightly excavated above hind coxae,
posteriorly with blunt apical angle. Metathorax long, convex [ventrally].
Legs with rather short forefemora, slightly thickening; [fore-] tibiae also
short, straight; [fore-] tarsi not shorter than tibiae, narrow. Hind femora not
shorter than forefemora, slightly thickening; [hind] tibiae straight. Hind tarsi
and middle legs not preserved. Unguiculate segment of foretarsi short and
almost triangular.

Dimensions. Body length 3.4 mm, length of rostrum 0.8 mm.

Comparison. Distinguished by position of rostrum, distally shifted place
of antennal attachment, and pronotum with distinct anterior angles.

Remarks. If the position of the rostrum is not the result of distortion
during fossilization, then this species should be considered independent of
other Jurassic rynchophoran beetles, and as transitional to forms having the
rostrum situated medially on the head capsule. The marked apical shifting of
the antennal attachment on the rostrum is also noteworthy. A slightly shifted
position of the rostrum is noted in O. similis described above. Oxycorynoides
progressivus probably belongs to an independent genus, standing somewhat
closer to forms with rhynchitoid rostrums than implied by its specific name.

Genus Scelocamptus L. Arnoldi, gen. nov.

Genus name coined from ‘skelis’ (Greek)—leg, and ‘komptos’ (Greek)—
curved.

Type species. S. tenuirostris, sp. nov.; Upper Jurassic of South Kaza-
khstan.

Diagnosis. Small, convex bettles. In lateral view of specimen, rostrum
typically belidoid, i.e., originating from lower part of head capsule, frontal
surface forming an angle of 95—120° with upper surface of rostrum. Eyes
small or large, rostrum smoothly curved, thinning toward apex, antennae

*The discussion at the beginning of the remarks section, and the use of the feminine pronoun
in the Russian original, imply that ‘it’ refers to the rostrum and not the entire specimen—Scien-
tific Editor.

224

attached almost midrostrum, their first segment not elongate. Pronotum
rather long. Elytra convex. Legs with moderately or markedly thickened
femora and always with curved tibiae, particularly fore - and hind tibiae; tarsi
small.

Species composition. Three species in the Late Jurassic of South
Kazakhstan.

Comparison. Distinguished by fore- and hind tibiae strongly curved in
apical third.

Scelocamptus tenuirostris L. Arnol’di, sp. nov.
(Plate XIII, Photo 7; Figure 96)

Species name coined from ‘tenuis’ (Latin)—slender, and ‘rostrum’ (Latin)—
beak.

Holotype. No. 2554/732, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailo-
vka site), Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head rather large, with slightly tuberculate frontal surface;
eyes large, placed at sides of head capsule, closer to lower margin. Rostrum
slender, rather long, but noticeably shorter than head capsule and pronotum
together; tapering toward apex. Antennae attached midrostrum, their first two
segments equal in length. Pronotum comparatively large, weakly emarginate
anteriorly, middle of base with two angular projections evidently positioned
against scutellum, which is not visible. Punctation noticeable on disk. Elytra
more than three times as long as pronotum, with sharp apical angle. Lateral

Fig. 96. Scelocamptus tenuirostris, sp. nov.; holotype PIN No. 2554 / 732;
Karatau, Upper Jurassic.

re ee te Set oh

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225

margin strongly emarginate above hind coxae. [Ventral] Structure of thorax
and abdomen indistinct; but, metathorax undoubtedly long and convex.
Structure of abdomen unclear. Legs with moderately thickened femora,
middle femora not thicker than hind ones; fore- and hind tibiae sharply curved
in apical third (middle legs not preserved). Tarsi short, not thick.

Dimensions. Body length 3.6 mm, length of rostrum 1.1 mm.

Comparison. Distinguished by moderately thickened femora, fore-
femora not thicker than hind ones.

Scelocamptus curvipes L. Arnol’di, sp. nov.
(Figure 97)

Species name coined from ‘cuvus’ (Latin)—curved, and ‘pes’ (Latin)—foot.

Holotype. No. 2384/512, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Holotype. The beetle undoubtedly strongly distorted during
fossilization and markedly flattened. Hence, all the less sclerotized parts are
stretched out along the vertical axis, particularly in anterior half.

Description. Head large, high,* frontal surface forming an angle of
approximately 100° with upper surface of rostrum. Eyes large, round, placed
higher than upper side of rostral base. Rostrum short, thick, noticeably
curved, gradually thinning toward apex. Antennae attached at end of basal
third of rostrum. Their structure indistinct. Pronotum large, uniformly

Fig. 97. Scelocamptus curvipes, sp. nov.; holotype PIN No. 2384/512; Karatau,
Upper Jurassic.

*See note at bottom of page 208—Scientific Editor.

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226

moderately convex, anterior margin weakly emarginate. Elytra dorsally very
gently convex, rounded apically, epipleural margin scarcely emarginate
above hind coxae. Metathorax long, weakly convex [ventrally]; abdomen
with similar sternites. Legs with strongly thickened forefemora, their width
more than one-third the length. Middle and hind femora noticeably thinner,
also hind femora shorter than forefemora. Fore- and hind tibiae rather
strongly curved along their entire length, middle tibiae very slightly curved;
tarsi small, slender.

Dimensions. Body length 4.0 mm, length of rostrum 1.3 mm.

Comparison. Distinguished by strongly thickened femora, hind femora
shorter and thinner than forefemora.

Scelocamptus dubius L. Arnol’di, sp. nov.
(Figure 98)

Species name coined from ‘dubius’ (Latin)—doubtful.

Holotype. No. 2066/3037, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley. Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Head small, diameter of eyes almost equal to basal width
of rostrum; frontal surface of head capsule hardly visible from above,
indistinct as it steeply descends to the upper surface of rostrum. Rostrum
slightly thicker than distance between eyes, considerably shorter than head
capsule and pronotum together, slightly broadening apically. Owing to the
semi-lateral position of the specimen, it is not clear whether the rostrum is
curved, as in the preceding forms, or more or less straight. Antennae attached

Fig. 98. Scelocamptus dubius, sp. nov.; holotype PIN No. 2066/3037; Karatau,
Upper Jurassic.

16

~

227

almost midrostrum, length of first four segments equal. Pronotum slightly
convex, with weakly projecting sides, widest in front of base, almost straight,
anterior margin slightly angularly projecting. Elytra long, ovoid, with some-
what round presutural angle, disk with distinct punctate grooves. [Ventral
surface of] thorax and abdomen not visible. Length of elytra 2.5 times that
of pronotum. Legs with slender but not long femora. Fore- and hind tibiae
noticeably curved along entire length; tarsi small.

Dimensions. Body length 4.6 mm, length of rostrum 1.8 mm.

Comparison. Distinguished by more slender femora.

Remarks. In general appearance this species most closely resembles
members of the preceding genus, but its tibiae are distinctly curved, a
fundamental feature of this genus.

Genus Ampliceps L. Arnoldi, gen. nov.

Genus name coined from ‘amplus’ (Latin)—large, and ‘caput’ (Latin)—
head.

Type species. A. dentitibia, sp. nov. Upper Jurassic of South Kaza-
khstan.

Diagnosis. Minute beetles, head highly convex frontally, forming less
than a right angle with upper side of rostrum, its gular surface coplanar with
lower side of rostrum. The latter rather thick and long, very slightly curved,
thinning toward apex. Eyes small, placed higher than level of rostral attach-
ment. Pronotum rather large, dorsally more or less flat. Elytra relatively
short, with acute-angled presutural apices. Legs short, femora distinctly
clavate; tibiae straight, short, with apical spurs.

Species composition. Two species in the Late Jurassic of South Kaza-
khstan.

Comparison. Distinguished by distinctly thickened femora and straight
tibiae.

Ampliceps dentitibia L. Arnol’di, sp. nov.
(Plate XIV, Photo 1; Figure 99)

Species name coined from ‘dens’ (Latin)—tooth, and ‘tibia’ (Latin)—shin.

Holotype. No. 2239/1562, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Description. Head large, frontal surface forming somewhat less than
right angle with rostrum, strongly convex. Eyes small, considerably less than
rostral width. Rostrum long, almost as long as head capsule and pronotum
together, uniformly tapering toward apex, weakly and uniformly curved.

228

Fig. 99. Ampliceps dentitibia, sp. nov.; holotype PIN No. 2239/1562; Karatau,
Upper Jurassic.

Place of attachment and shape of antennae not established. Pronotum with
smooth anterior margin and rounded base, finely and densely punctate on
disk. Elytra broad, with slightly curved epipleural margin and apices right-
angled. Metathorax long, convex [ventrally]. Abdomen with similar ster-
nites, pygidium present. Legs with short, moderately clavate femora,
forefemora slightly longer than others, tibiae almost straight, a pair of spurs
on inner angle of foretibiae, and apparently also on other tibiae but somewhat
shorter. Tarsi slender, short, their first segment longer than remaining seg-
ments, unguiculate segment almost equal to it in length.

Dimensions. Body length 2.5 mm, length of rostrum 1.0 mm.

Comparison. Distinguished by longer head, and paired spurs on inner
angle of foretibiae.

Ampliceps furcitibia L. Arnol’di, sp. nov.
(Figure 100)

Species name coined from ‘furca’ (Latin)—fork, and ‘tibia’ (Latin)—shin.

Holotype. No. 2239/1551, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype. m

Description. Head high , but short; eyes small, less than basal width of
rostrum; frontal surface of head capsule convex, forming an angle of about
80° with upper side of rostrum. Rostrum rather thick, slightly tapering toward
apex, almost straight. Eyes placed above rostrum. Place of attachment and
shape of antennae not clear. Anterior margin of pronotum slightly emar-

*See our note on page 208—Scientific Editor.

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229

ginate, base of pronotum very weakly excavated; disk with dense, moderately
large punctures. Elytra comparatively long, noticeably tapering toward apex,
2.2 times ionger than pronotum, their apical angles acute. Epipleural margin
weakly excavated. [Ventral surface of] thorax and abdomen not visible. Legs
with short, moderately clavate femora; forefemora thickest; tibiae straight;
foretibiae longer than others, apically appearing bifurcate for a short distance
with spurs on outer and inner angles. Foretarsi slender, their first segment
larger than remaining segments. Middle and hind tarsi not preserved.

Dimensions. Body length 4.5 mm, length of rostrum 1.5 mm.

Comparison. Distinguished by high [= tall] but shori head, and two spurs
on opposite sides of tibial apex.

Genus Paroxycorynoides L. Arnol’di, gen. nov.

Genus name coined from genus Oxycorynoides.

Type species. P. elegans, sp. nov.; Upper Jurassic of South Kazakhstan.

Diagnosis. Body elongate. Head small, rostrum slightly tapering from
base to midrosiral antennal attachment, then broadening toward apex, api-
cally slightly flared angularly. Mandibles small. Antennae with very slightly
elongate first segment and non-abrupt three-segmented club, posteriorly
almost extending to middle of pronotum. Pronotum transverse, with rather
deep anterior emargination, sides uniformly rounded, base excavated. Elytra
elongate, almost 3.5 times longer than pronotum, with acute apical angles.

Species composition. Monotypic genus.

Comparison. Differs from other representatives of the tribe in long and
narrow body; unique rostrum; small, short pronotum, and long elytra.

Fig. 100. Ampliceps furcitibia, sp. nov.; holotype PIN No. 2239/1551; Karatau,
Upper Jurassic.

230

Remarks. Owing to the position of specimen on its ventral side, a
detailed comparison with other representatives of the tribe becomes difficult.
Hence, its inclusion in this tribe is not verified.

Paroxycorynoides elegans L. Arnol’di, sp. nov.
(Plate XIV, Photo 2; Figure 101)

Species name coined from ‘elegans’ (Latin)—elegant.

Holotype. No. 2452/125, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Uspenskoe (Karatau-Galkino site). Upper Jurassic,
Karabastau series.

Material. Holotype.

Description. Head small, round. Eyes small, less than basal width of
rostrum shifted forward. Rostrum noticeably longer than head capsule and
pronotum together. Pronotum round, tapering anteriorly, its posterior margin
1.8 times the anterior, maximum width of pronotum 1.7 times its medial
length. Scutellum easily discernible, almost pentagonal. Elytra 1.7 times
longer than maximum width. Abdomen projecting a little beyond apex of

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Fig. 101. Paroxycorynoides elegans, sp. nov.; holotype PIN No. 2452/125;
Karatau, Upper Jurassic.

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231

elytra. Legs, judging from the faint impression of part of middle femur, not
thick.
Dimensions. Body length 3.1 mm, length of rostrum 1.1 mm.

Tribe ECCOPTARTHRINI L. Arnol’di, trib. nov.

Diagnosis. Small beetles. Rostrum rather long, antennae attached close to its
middle. Pronotum transverse. Elytra broad. Legs short, femora thickened,
first tarsal segment much larger than remaining segments, broad, almost
bilobed, notched apically.

Composition. Tribe includes a single genus.

Comparison. Distinguished by completely abnormal form of foretarsi
with very large, almost bilobed first segment.

Remarks. Enlargement of first tarsal segment also noted in some Jurassic
belidoid Rhynchophora, but its structure in the latter is totally different.

Genus Eccoptarthrus L. Amoldi, gen. nov.

Genus name coined from ‘eccoptio’ (Latin)—notched, and ‘arthron’
(Greek)—joint.

Type species. E. crassipes, sp. nov.; Upper Jurassic of South Kaza-
khstan.

Diagnosis. Head rather short. Antennae with small club. Pronotum very
weakly transverse. Foretibiae curved.

Eccoptarthrus crassipes L. Arnoldi, sp. nov.
(Plate XIV, Photo 3; Figure 102)

Species name coined from ‘crassus’ (Latin)—thick, and ‘pes’ (Latin)—foot.

Holotype. No. 2239/1507, PIN, almost entire impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank
of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Eyes shifted forward, frons between them almost the same
width as base of rostrum. Rostrum preserved only in basal half, apparently
rather long, place of antennal attachment close to midrostrum. Antennae only
partially preserved, and had non-abrupt club. Punctation visible on head
capsule. Maximum width of pronotum 1.2 times its length, its anterior margin
rather deeply emarginate, posterior projecting backward at obtuse angle,
laterally almost parallel, with rather coarse punctation. Elytra broad, with
rows of punctures in grooves. Triangular scutellum visible between elytral
bases. Legs with short, thick femora, forefemora thicker than remaining

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232

Fig. 102. Eccoptarthrus crassipes, sp. nov.; holotype PIN No. 2239/1507;
Karatau, Upper Jurassic.

femora; foretibiae curved basally, straight further on middle tbiae slightly
shorter, straight. First tarsal segment longer than the two following together.
Dimensions. Body length about 6 mm, rostrum longer than 1.5 mm.

Tribe DISTENORRHININI L. Arnol’di, trib. nov.

Diagnosis. Rostrum moderately long, noticeably constricted at place of
antennal attachment, again constricted preapically or before base. Place of
antennal attachment in anterior third or even anterior fourth of rostrum. Apex
of latter slightly broadening, mandibles of usual size. Pronotum transverse.
Elytra elongate, flat. Legs with poorly broadened femora.

Composition. Tribe includes single genus.

Comparison. Distinguished by unique structure of rostrum having two
constrictions.

Genus Distenorrhinus L. Arnoldi, gen. nov.

Genus name coined from ‘di’ (Greek)—two, ‘stenos’ (Greek)—narrow, and
‘rhinos’ (Greek)—nose.

Type species. D. angulatus , sp. nov.; Upper Jurassic of South Kaza-
khstan.

233

Diagnosis. Head transverse. Rostrum almost as long as head and
pronotum together. Pronotum transverse. Femora slightly thickened, tibiae
straight, slender.

Species composition. Two species in the Late Jurassic of South
Kazakhstan.

Remarks. A comparison of the two species in this genus shows sig-
nificant differences in their rostral structure, and possibly their inclusion
under one genus is provisional. Distenorrhinus has some features in common
with the representative of the next subfamily, Brenthorrhininae, particularly
the structure of the antennae and their nearly apical attachment.

Distenorrhinus angulatus L. Amol’di, sp. nov.
(Plate XIV, Photo 4; Figure 103)

Species name coined from ‘angulatus’ (Latin)—angular.
Holotype. No. 2239/1547, PIN, almost enure impression of beetle.
South Kazakhstan, Chimkent oblast, Algabass region, south-western flank

Fig. 103. Distenorrhinus angulatus, sp. nov.; holotype PIN No. 2239/1547;
Karatau, Upper Jurassic.

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234

of Kashkarata river valley, Aulie area near Mikhailovka village (Karatau-
Mikhailovka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Eyes strongly shifted forward. They were probably marke-
dly convex. Rostrum almost as long as pronotum and head capsule together,
tapering initially sharply, then gradually to the place of antennal attachment
at the end of distal third of rostrum. Constriction rather sharp at antennal
attachment, from which point for a short distance the rostrum again broadens,
then again narrows. Second constriction somewhat weaker than first, apical
expansion weak. Antennae poorly preserved, but their first segments ob-
viously small. Pronotum weakly transverse, its anterior angles pointed and
directed somewhat laterally; sides slightly rounded, anterior margin with
shallow emargination, posterior margin straight, slightly narrower than
anterior. Elytra elongate, each three times longer than its maximum width,
for two-thirds of length almost parallel-sided, with rounded apices, punctate
grooves easily discernible, punctures small. Metathorax and abdomen not
preserved. Legs slender, slightly thickening in the middle.

Dimensions. Body length 3.8 mm, length of rostrum 1.3 mm.

Comparison. Distinguished by the second rostral constriction more
distal to the place of antennal attachment.

Distenorrhinus antennatus L. Arnol’di, sp. nov.
(Figure 104)

Species name coined from ‘antenna’ (Latin)—feeler.

Holotype. No. 2554/721, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-
ka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Length of head capsule half its width at pronotal margin;
eyes small, shifted forward; anterior margin of head capsule well differen-
tiated, frontally steeply descending to rostral base. Rostrum noticeably longer
than head capsule and pronotum together. Distance between anterior margins
of eyes 1.6 times the thickness of rostrum. For a distance of one-fourth its
length from base, rostrum briefly and shallowly constricted, then reattaining
its former width, and further for about half its length gradually narrowing to
the place of antennal attachment at the commencement cf its distal fourth.
Then rather noticeably broadening to apex. Apex of rostrum slightly broader
than its base. Mandibles normal in size. Antennae long, their apices extending
beyond middle of pronotum; first antennal segment one-and-a-half times
longer than second; third twice the length of first; remaining segments
elongate, scarcely shorter than first; club distinct, its last segment

ee

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235

Fig. 104. Distenorrhinus antennatus, sp. nov.; holotype PIN No. 2554/721;
Karatau, Upper Jurassic.

rhomboidal. Pronotum strongly transverse, its width 1.7 times its medial
length, smoothly emarginate anteriorly and posteriorly, sides uniformly
shallowly emarginate, undoubtedly flattened dorsoventrally. Elytra elongate,
their length 3.3 times their maximum width, apically rounded. Metathorax
long. Legs moderately long, femora weakly clavate.

Dimensions. Body length 4.0 mm, length of rostrum 1.3 mm.

Comparison. Distinguished by proximal position of second rostral con-
striction.

Subfamily Brenthorrhininae L. Arnol’di, Subfam. Nov.

Diagnosis. Rostrum short, thick, strongly broadening apically. Antennae
attached at apex of rostrum. Mandibles large. Pronotum strongly transverse.
Elytra parallel-sided, abdominal sternites similar.

Composition. Single genus.

Comparison. Distinguished by apical attachment of antennae.

Remarks. There is only one representative of this subfamily in the
material studied by the author. Due to the impression being positioned on its
dorsal surface, it is not possible to reliably establish the angle between the
rostrum and head capsule (i.e., whether belidoid or not) nor whether distinct
noto-pleural ridges were present on the pronotum. In general appearance the

236

beetle somewhat resembles brentids. The structure of its rostral apex very
closely resembles that observed in males of Recent brentids. The antennal
structure has no correspondence in Recent species of the family, and is
absolutely unique.

Genus Brenthorrhinus L. Armol’di, gen. nov.

Genus name coined from genus Brenthis, and ‘rhinos’ (Greek)—nose.
Type species. B. mirabilis, sp. nov. Upper Jurassic of South Kazakhstan.
Diagnosis. Head large, transverse; rostrum noticeably shorter than head

capsule and pronotum together, thick, sharply broadening apically, its thick-

ness equal to one-fourth its length. Antennae attached at very corners of
rostrum, their first segment strongly arched and only slightly shorter than
apical width of rostrum, second one-third, the length third straight and as
long as first, remaining segments twice as long as wide, club non-abrupt but
distinct. Pronotum dilated, anteriorly rather strongly emarginate, with dis-
tinct ridge, i.e., having narrow constriction. Elytra parallel-sided, with right-
angled apices. Metathorax long, three times the length of hind coxae.

Abdomen with similar sternites. Legs long, femora rather strongly thickened,

forefemora larger than others; tibiae thin, very slightly curved in anterior half,

with short apical bristles.
Species composition. Monotypic genus.

Fig. 105. Brenthorrhinus mirabilis, sp. nov.; holotype PIN No. 2554/708: a—dor-
sal view; b—ventral view; Karatau, Upper Jurassic.

173

23%)

Brenthorrhinus mirabilis L. Arnoldi, sp. nov.
(Plate XIV, Photo 5; Figure 105)

Species name coined from ‘mirabilis’ (Latin)—remarkable.

Holotype. No. 2554/708, PIN, almost entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley, Aulie area near Mikhailovka village (Karatau-Mikhailov-

.ka site). Upper Jurassic, Karabastau series.

Material. Holotype.

Description. Mandibles only slightly shorter than apical width of
rostrum. Mentum visible between them, anteriorly extending up to almost
half the length of mandibles. Head capsule laterally transforming stepwise
into rostrum. Third antennal segment reaching [back to] this step, whence
the antennae, instead of running parallel to rostrum, turn at an angle as in
geniculate antennae of Recent forms, though in this specimen they are
certainly not geniculate. Prosternum with broad process between forecoxae,
posterior angles completely rounded. Abdomen slightly shorter than large
metathorax. Elytra with slender punctate grooves.

Dimensions. Body length 6.7 mm, width at shoulders 2.6 mm, length of
rostrum 1.7 mm.

Subfamily Nanophydinae L. Arnoldi, Subfam. Nov.

Diagnosis. Rostrum slightly curved, with antennae attached at middle.
Metathorax very short. Fourth abdominal sternite with posterior angular
notch.

Composition. Single genus.

Comparison. Distinguished by characteristic structure of abdomen with
angularly notched fourth sternite, and short metathorax.

Genus Nanophydes L. Arnoldi, gen. nov.

Genus name coined from genus Nanophyes.

Type species. N. ovatus, sp. nov. Upper Jurassic of South Kazakhstan.

Type species. Body small, shortand wide, its general configuration oval.
Head large. Rostrum rather long, curved. Pronotum short. Metathorax ap-
proximately one-third of abdomen in length. First three abdominal sternites
of normal structure, fourth short, posteriorly with wide deep emargination
into which obtuse angle of fifth sternite fits.

Species composition. Monotypic genus.

Remarks. The shortened metathorax and shape of body strongly suggest
that this form had lost the ability to fly.

238

Fig. 106. Nanophydes ovatus, sp. nov.; holotype PIN No. 1789/102; Karatau,
Upper Jurassic.

Nanophydes ovatus L. Arnol’di, sp. nov.
(Plate XIV, Photo 6; Figure 106)

Species name coined from ‘ovum’ (Latin)—egg.

Holotype. No. 1789/102, PIN, entire impression of beetle. South
Kazakhstan, Chimkent oblast, Algabass region, south-western flank of Kash-
karata river valley. Uspenskoe village (Karatau-Calkino site). Upper Juras-
sic, Karabastau series.

Material. Holotype.

Description. Head large; eyes small, placed above upper side of rostrum.
Length of rostrum noticeably exceeding length of head capsule and pronotum
tegether; rostrum strongly but gradually tapering toward apex, and gently
curved all along entire length; antennal attachment close to midde. Pronotum
short, with large sparse punctures, posterior angles rounded. Judging from

174 the shape of metathorax and abdomen, the elytra were short and wide, with
distinct epipleura. Legs comparatively short, with slightly clavate femora,
and apparently straight, slender tibiae.

Dimensions. Body length 3.0 mm, length of rostrum 1.6 mm.

* *e *

The author acknowledges the fact that the classification of Eobelidae
followed above is largely tentative and partly artificial. The main reason for
this is the nature of the material itself. The fossil beetles studied in this work
represent a more or less random selection from the entire fauna. Moreover,
this material did noi fossilize simultaneously but over some period of time
that was extremely long from the viewpoint of an entomologist studying
extant fauna. Another point of great significance is that generally it is not
possible to study the entire repertoire of taxonomic characters, but only those
which are visible in an impression.

239

However, the persistence of the main structural features of all the
Jurassic Rhynchophora studied, excluding a few doubtful cases not differing
from those of the above described primitive type, cannot be incidental. This
certainly reflects a definite evolutionary level which this already highly
specialized group of Coleoptera had achieved.

The overall primitiveness of Eobelid morphology has already been
established; this family combines within it the most primitive characters
which are individually preserved in various extant groups of Rhynchophora
presently having the status of independent families. The individual characters
[taken in combination] are peculiar to the fossil forms. [The preceding
observations] definitely support the idea of a common origin for what is
currently an enormous group. Based on general appearance and most of the
main characters, the Late Jurassic Rhynchophora most closely resemble the
primitive representatives of the Recent family Belidae. Therefore, they may
tentatively be considered the “beloidoid’* phase of rhynchophoran evolu-
tion.

Among the Late Jurassic Rhynchophora, the most stable morphological
structure is the rostrum. This structure, the main diagnostic feature of the
group, arose much earlier and reached a high level of specialization in the
Late Jurassic. The author suggests that the development of the rostrum
assisted the group in becoming the largest of all in the order Coleoptera, since
it made it possible for the preimaginal stages to develop within the plant
tissues or under their cover. This provided the larvae with an abundance of
high-quality food, more or less constant optimal atmospheric humidity and
probably a significant reduction in the pressure of predators and parasites. In
the Late Jurassic forms studied, the rostra are of the same type with a solitary
exception, and are closest in structure to those of Recent Nemonychidae or
Rhynchitinae of the family Attelabidac. The rostrum of Brenthorrhinus
differs most significantly from this type: it is short and thick, with an apical
attachment of the antennae, a feature now more characteristic of species with
terrestrial larvae. However, its structure is very similar to that in males of
some Brentidae, and is probably a secondary sexual character as is true for
the highly enlarged mandibles. In detail, the rostrum of the studied Late
Jurassic forms has arather varied structure and probably reflects the frequent
species or group adaptations to different plant species or different plant parts.
It must be emphasized again that the studied examples of Late Jurassic
Rhynchophora have a lower placement of the rostrum, which is presently
characteristic only of the relictual family Belidae. All the remaining Recent
families have a middle or upper position of the rostrum. The middle position
is characteristic of the most morphologically primitive families: Nemo-

*So given in the Russian original. In the context of its reference to the family Belidae it would
be more correct to say “belidoid’”—General Editor.

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240

nychidae, Oxycorynidae, Attelabidae (subfamily Rhynchidae), and a few
others. All of the extant Curculionidae have an upper position of the rostrum.

Among the Recent forms, the notopleural ridge has only been preserved
in a few Oxycorynidae (also a relict family). In a few Curculionidae a faint
blunt ridge is present laterally on the pronotum, but there is never a sharp
flange and notopleural suture. It is interesting to note that the notopleural
ridge and suture are absent in the extant Belidae.

The antennae of Eobelidae are closest to those of the Recent Attelabidae
(subfamily Rhynchitinac) or to Nemonychidae. They always have a loose
and indistinct three-segmented club which is absent in the extant Belidae.

The aforementioned facts strongly suggest that the Mesozoic Rhyn-
chophora of the “Belidoid” type (having a low position of the rostrum) later
became extinct to a large extent, leaving behind a dead-end, relictual group:
the extant Belidae. It is hardly possible to trace direct descendants of the
already specialized Jurassic Rhynchophora among the Recent cur-
culionids.The Late Jurassic rhynchophoran fauna, apparently rich in species,
fed on different plants of their period and had diverse trophic relationships
which are reflected in the varied types of rostra, legs, and general body
shapes. However, judging from the present-day associations of curculionids,
it may be assumed that their trophic rejationships with Cryptogams were
never widespread, and the rare modern occurrences represent a secondary
adaptation.

Several general conclusions pertaining to the further evolution of the
Rhynchophora can be drawn from the study of the available material. Based
on the general structure of the elytra and their deformation during fossiliza-
tion, it may be assumed that the elytra of the Jurassic Rhynchophora were
relatively weakly sclerotized and mostly flattened. More durable structures
were the prothorax, mesothorax, metathorax, and head, particularly the
rostrum. The subsequent independent evolution of all representatives of the
group proceeded in parallel toward a greater degrce of sclerotization, greater
convexity of the elytra and prothorax, and disappearance of the notopleural
ridge. The abdomen of most of the Rhynchophora lost the homonomic
structure of the sternites, all of which, except the first, were evidently
movable. In the vast majority of extant Rhynchophora, the first two sternites
are partially or completely fused and occupy more than half the length of the
abdomen; mobility has been retained only in the abdominal apex. The fusion
of many sutures in the thoracic region proceeded in parallel. It may be
speculated that all these modifications occurred under the influence of
general climatic aridity, as well as the increasing pressure of entomophages.

The position of the antennal attachment was markedly different in most
of the Jurassic species examined. It was near the midrostrum, rarely close to
its apex, but never at its base. In the extant belids and oxycorynids, on the
contrary, the antennae are always attached to the base of the rostrum or in its

Ee ope He ae

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241

proximal third (Belidae). The disappearance of the antennal club is note-
worthy in the Recent Belidae, the Jurassic ancestors of which always had a
distinct (though not abrupt), loose club of the “rhynchitoid” type. Based on
the absolute predominance of a well developed club among the extant
Rhynchophora, this structure can be considered advanced; the reasons for its
reduction in the Recent Belidae are unclear.

As previously mentioned, the belidoid type is distinctly predominant
among the studied examples of Jurassic Rhynchophora. However, several
forms, while preserving the fundamental archaic belidoid characters, are
markedly similar to the most primitive of the extant oxycorynids, particularly
in the structure of the pronotum and the general body shape. More rarely,
they are similar to the Recent Nemonychidae and Rhynchitinae of the family
Attelabidae. None of the aforementioned groups, which deviated from the
generalized belidoid type, can be considered as a main evolutionary line
equal in importance to the latter. Judging from the available material, these
forms which deviated from the belidoid group never flourished, and therefore
cannot be rated as its “sister” group. Thus, within the [deviant] group, several
evolutionary branches arose simultaneously. These may be regarded as a
“cluster” of new lines of specialization, and not [the result of] a dichotomous
branching [which would have yielded] a new sister group (sensu Hennig,
1969) of equivalent status [to the generalized belidoid line].

It should be emphasized again that all the descendant groups of the
Rhynchophora complex that flourished in the Late Jurassic are now repre-
sented only by small relictual families. There is as yet not adequate data for
establishing the time of separation from the “belidoid” stock of the forms
which could be considered representatives of the extant Curculionidae.

In his interesting review of paleogeographic data on insects, Zherikhin
(1970) prefers to group the Late Jurassic Rhynchophora with the family
Oxycorynidae and only to a lesser degree with Belidae. However, many of
the abovenoted structural features of the Jurassic species, which are most
important to explain the taxonomic affinity of these insects, force us to
consider them as representatives of a separate family, now extinct, which is
closest to the Recent Belidae but has several characters of the family
Oxycorynidae as well as Nemonychidae and Attelabidae.

Plesiomorphic characters have never been observed simultaneously in
a more or less complete “selection” of the Recent relict Rhynchophora.
However, a set of the more important ones (for example, the structure of the
head capsule and rostrum, or the structure of pronotum, etc.) characterizes
each of the extant families. On the other hand, within a complete range of
the Late Jurassic Rhynchophora all the plesiomorphic characters are usually
simultaneously present, a reaffirmation of the monophyletic origin of the
Rhynchophora.

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Family ATTELABIDAE Billberg, 1820
Subfamily Rhynchitinae Thomson, 1859
Tribe AULETINI Reitter, 1912
Genus Baissorhynchus Zherichin, gen. nov.

Genus name coined from Baisa river, and ‘rhynchos’ (Greek)—rostrum.
Type species. B. tarsalis, sp. nov. Lower Cretaceous of Trans-Baikal.
Description. Head short, hemispherical, transverse, lacking constriction

beyond eyes. Temples short. Eyes large, round, lateral. Gular suture ap-

parently medial. Rostrum long, rather slender, cylindrical, uniformly and
rather strongly curved, with transverse depression at junction with head.

Labrum labular, free. Prosternum without traces of sutures. Elytra slightly

convex, with steep, almost vertical slope, covering pygidium, with irregular

rows of punctures. Epipleura reduced, but distinct. External basal angle of
elytra oblique. Wings normally developed. Forecoxae conical, approaching
posterior margin of prosternum. Middle coxae spherical, contiguous. Femora
short, rather broad, lacking armature. Tibiae same length as femora. Tarsi
with bilobed, deeply cleft second and third segments; true fourth segment
not visible.

Species composition. One species in Early Cretaceous of Trans-Baikal.

Comparison. Differs from all the known genera of the tribe in the deeply
cleft, bilobed second tarsal segment, similar in shape to third. [Also differ in
possessing] a free labrum, short temples, and irregular rows of punctures on
elytra instead of usual scattered punctation. Scattered punctation is present
on earlier described genera with the exception of Car Blackb. from which

Baissorhynchus is distinguished by strongly curved rostrum, and consider-

ably shorter and thicker legs as well as a [different] structure of tarsi.
Remarks. The state of preservation of the holotype Baissorhynchus

tarsalis Zherichin sp. nov., does not permit the clarification of certain
taxonomically important structural details. Among the characters available
for study, the structure of labrum is very important. A free labrum is present
only in a few Curculionoidea, viz., Rhinomaceridae, Anthribidae, Platy-
podidae, some Attelabidae (taking the latter family in the sense suggested by
R. Crowson, 1955), and also in an anomalous genus Anchylorrhynchus Fahr.
from the family Curculionidae (Voss, 1943). A long rostrum is never ob-
served in the families Anthribidae and Platypodidae. The presence of distinct
elytral epipleura argues against the inclusion of Baissorhynchus in Cur-
culionidae. It must therefore be included in either Rhinomaceridae or At-
telabidae which it even resembles in general appearance. The main
distinguishing characters by which one can reliably separate representatives
of these two families (structure of mandibles and maxillary palps) cannot be
established from the available remains or the wing venation. The nature of

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243

the gular suture supports the inclusion of Baissorhynchusin Attelabidae. This
suture is paired in ail the known Rhinomaceridae, whereas in Attelabidae
only one medial suture is present. There is no trace of the paired gular suture
on the medial cleavage line on the head of the Baissorhynchus tarsalis
holotype; it is assumed that it was medial and proceeded along the cleavage
line.

Judging from the shape of the head and rostrum, Baissorhynchus must
be included in the subfamily Rhynchitinae of the family Attelabidae. Com-
parative characters draw it close to the isolated genus Car Blackb. (four
contemporary species from Australia, and one from the Paleogene of
Europe). Car also exhibits several other characters distinguishing it from the
remaining Rhynchitinae—such as simple free claws, basally attached anten-
nae and free abdominal sternites—but unfortunately it is not known whether
Baissorhynchus also possessed them. Since the genus Car is not known to
us in nature and its comprehensive comparison with Baissorhynchus is out
of question, it seems reasonable to postpone separating these general into a
special taxon and retain them in the tribe Auletini of the subfamily
Rhynchitinae, where Car was placed by Crowson (1955). It is noteworthy
that the structure of the tarsi in Baissorhynchus is different from that in
Attelabidae and Rhinomaceridae, but resembles that in oxycorynids.

Baissorhynchus tarsalis Zherichin, sp. nov.
(Plate XIV, Photo 7; Figure 107)

Species name coined from ‘tarsus’ (Latin)—foot.

Holotype. No. 1989/3010, PIN, reverse impression of almost entire
beetle. Trans-Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river,
downstream from the mouth of Baisa river (Baisa site). Lower Cretaceous,
Zazin series.

Material. Holotype.

Description. Rostrum equal in length to head and pronotum together,
with very fine dorsal rugae basally. Punctation on head fine, rather dense.
Pronotum not longer than its basal width, apparently faintly bi-emarginate,
dorsally almost flat, steeply sloping anterioriy just before anterior margin.
Punctation of pronotum large and dense; on sides almost partially fused into
transverse rugae. Punctures on elytra large, dense. Distance between punctate
rows not exceeding diameter of punctures. Tibiae straight. Tarsi slightly
shorter than tibiae, their first segment broad, second and third equal to it in
length, unguiculate segment almost equal in length to remaining segments
together. Sides of thorax and at least base of abdomen with dense coarse
punctures.

Dimensions. Body length (without rostrum) 2.2 mm.

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244

na
-
ae ==
Fig. 107. Baissorhynchus tarsalis, sp. nov.; holotype PIN No. 1989/3010; Baisa,
Lower Cretaceous.

Family CURCULIONIDAE Latreille, 1802

Subfamily Nanophyinae Seidlitz, 1891
Genus Cretonanophyes Zherichin, gen. nov.

Genus name coined from ‘creta’ (Latin)—chalk, and genus Nanophyes.

Type species. Cretonanophyes longirostris, sp. nov. Lower Cretaceous
of Trans-Baikal.

Description. Rostrum very long, delineated from head by a depression*.
Eyes large, round. Frons rather wide, without tubercle. Antennae geniculate
with thick six (or possibly seven) segmented flagellum. Pronotum apparently
with narrow basal slope. Scutelium clearly visible. Elytra with well
developed humeral tubercles, without tubercles on disk or lateral flanges.
Forecoxae reaching anterior margin of prosternum. Trochanters large.
Femora without denticles. Tibiae longer than femora, slender, straight, not
apically broadening. Second tarsal segment slightly bilobed. Claws slender,
free, without denticles, equal in length. Third and fourth abdominal sternites
strongly truncate. Pygidium free.

Species composition. One species in Early Cretaceous of Trans-Baikal.

Comparison. Differs from ali the known genera of the subfamily in
broader frons; thickened flagellar segments of antennae, equal in width to
apex of scape; clearly identifiable scutellum, and bilobed second tarsal
segment. Among the present-day Nanophyinae, free claws are present only
in the genus Corimalia Gozis. However, the species of this genus usually
possess a four- or five-segemented antennal flagellum; if six-segmented then

*The term ‘depression’ as given here does not refer to a pit or groove, but rather to a drop-off
or down-step—Scientific Editor.

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245

the femora are always with denticles. Antennal flagella in Ctenomerus
Schoenh., Hexatmetus Marshl. and Diplophyes Marshl. also consist of six
segments, but in all representatives of these genera the claws are fused and
the femora have denticles. Cretonanophyes differs further from Hexatmetus
Marshl. in the developed humeral tubercles of the elytra. The genus
Nanomicrophyes Pic, having free claws, is related not to Nanophyinae but to
Tychiinae. In appearance, Cretonanophyes is most similar to species of the
extant genus Ctenomerus Schoenh. (tropical Africa and Asia).

Remarks. The combination of geniculate antennae together with en-
larged trochanters undoubtedly indicates the affiliation of Cretonanophyes
to the subfamily Nanophyinae, family Curculionidae. This is also confirmed
by the characteristic inequality of the abdominal segments.

Cretonanophyes longirostris Zherichin sp. nov.
(Plate XIV, Photo 8; Figure 108)

Holotype. No. 1668/1772, PIN, almost entire impression of beetle: Trans-
Baikal, Buryat ASSR, Eravnin region, left bank of Vitim river, downstream
from the mouth of Baisa river (Baisa site). Lower Cretaceous, Zazin series.

Material. Holotype.

Description. Head rounded, with fine, sparse punctures. Rostrum almost
twice (7 : 4) as long as head together with pronotum, slender, cylindrical,
weakly and uniformly curved, with sparse punctation basally, in apical part
smooth or possibly with microscopic punctures. Antennae attached some-
what behind midrostrum. Scape short, not reaching base of rostrum, slender,
thickening club-like toward apex, curved. First two flagellar segments iden-
tical, longer than wide; third segment shorter than the two preceding; fourth
longer than third; fifth and sixth identical, slightly broader than preceding
ones, equal in length and width. Pronotum apparently not longer than wide,
convex, greatest height posterior to middle, dorsally and laterally with sparse,
fine punctures. Elytra rather strongly and uniformly convex, with rows of
sparse, large punctures. Intervals between them wider than diameter of
punctures, apparently with weak transverse rugae. Legs with dense, slender,
rather long, oppressed pubescence. Tarsi shorter and wider than tibiae, their
second segment longer than wide, third shorter than second, unguiculate
segment shorter than second and third together.

Dimensions. Body length (without rostrum) 3.1 mm.

Subfamily Slonikinae Zherichin, Subfam. Nov.
Diagnosis. Rostrum long, slender, cuneately narrowing toward apex, apical-

ly pointed. Antennal sockets directed toward eyes. Antennae not geniculate,
with loose club, consisting of individual rounded segments, attached in distal

246

Fig. 108. Cretonanophyes longirostris, sp. nov.; holotype PIN No.1668/1772;
Baisa, Lower Cretaceous.

half of rostrum. Prothorax with distinct lateral slope, without a groove for
insertion of rostrum. Elytra with punctate grooves. Forecoxae attached at
posterior margin of prosternum, contiguous. Middle coxae convergent.
Metathorax equal in length to first abdominal sternite; Trochanters small. All
femora identical, not thickened. Tibiae with projecting outer apical angle.
Abdominal sternites slightly varying in length.

Composition. One genus in Early Cretaceous of Trans-Baikal.

Comparison. Differs from most subfamilies of Curculionidae in non-
geniculate antennae; from Apioninae (having similar antennae) in short
trochanters and loose club; from Brachycerinae in long rostrum; from
Rhynchaeninae in not thickened hind femora, rostrum notcurved under body,
and apical process on tibiae; from Cryptoderminae in antennal attachment

180 far from rostral base, loose club, and elongate linnear antennal grooves.

Remarks. Genus Slonik undoubtedly belongs to the family Curculioni-
dae and has several archaic distinguishing characters (nongeniculate anten-
nae, loose club not clearly demarcated from flagellum, and lateral slope of
prothorax). In appearance it resembles some Tychiinae (for example, genus
Tychius Cerm.), but the latter has no species with nongeniculate antennae;
moreover, the drawn out posterior corners of the middle abdominal sternites
are characteristic of them. Among the families with first antennal segment
short, Slonik slightly resembles only Cryptoderminae. This is a relict group
including one oriental and one African genus. However, Cryptoderminae is

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an advanced group, aside from [possessing] the aforementioned
plesiomorphs, and is a derivative of Rhynchophorinae; the reduction of the
first antennal segment is clearly secondary in Cryptoderminae.

Genus Slonik Zherichin, gen. nov.

Name of the genus is a Latinized Russian word.*

Type species. S. sibiricus, sp. nov. Lower Cretaceous of Trans-Baikal.

Description. Head spherical, ventrally strongly convex. Temples very
short. Frons wide, wider than diameter of eyes. Rostral junction with frons
without depression**. Antennae slender, rather long, reaching posterior
margin of eyes. Scutellum large, triangular. Elytra keel-shaped, tapering
posteriorly with distinct rows of punctures. Humeral tubercles well
developed. Lateral margins of elytra straight. Prothorax anteriorly with small
hood, separated by shallow groove. Posterior margin of abdominal sternites
straight. Fifth sternite longer than others. Pygidium completely covered by
elytra. Femora without armature, without apical constriction. Tibiae slender,
straight, gradually broadening toward apex.

Composition. One species in Early Cretaceous of Trans-Baikal.

Slonik sibiricus Zherichin, sp. nov.
(Plate XIV, Photo 9; Figure 109)

Species named after Siberia.

Holotype. No. 1989/2938, PIN, direct reverse impressions of entire body
of well-preserved beetle. Trans-Baikal, Buryat ASSR, Eravnin region, left
bank of Vitim river, downstream from the mouth of Baisa river (Baisa site).
Lower Cretaceous, Zazin series.

Material. Holotype.

Description. Head with dense, coarse punctures. Eyes elongate; rostrum
equal in length to head and pronotum together, strongly and uniformly
curved, with slender keel above antennal sockets. Punctation of rostrum
rather dense and coarse. Antennae attached at one-third length of rostrum
from its apex. Their first six segments almost identical, cylindrical, consid-
erably longer than wide. Seventh and eighth segments slightly broader and
considerably shorter, almost not longer than wide. Ninth rounded, broader
than eighth, but narrower and shorter than the following two. These last are
markedly enlarged, rounded. Pronotum flat, with straight posterior margin,
and dense, coarse punctation. Elytra flat, sloping from base to apex, with
rows of sparse large punctures. Intervals between them broader than diameter

**Slonik’ is a transliteration of the Russian word which means ‘little elephant’. This term is
often applied to weevils—Scientific Editor.
**See comment on bottom of p. 244—Scientific Editor.

180

248

Fig. 109. Slonik sibiricus, sp. nov.; holotype PIN No. 1989/2938; Baisa, Lower
Cretaceous.

of punctures. Prothorax with transverse rugose punctures. Punctation of
meso- and metathorax and abdomen dense and rather coarse. Body dark
colored, antennae and legs light colored, antennal clubs dark.

Dimensions. Body length (without rostrum) 3.1 mm.

REMARKS ON THE PHYLETIC LINKS AND ECOLOGY OF
BAISIAN RHYNCHOPHORA

Three families of Rhynchophora are represented in the Early Cretaceous
Baisian fauna: Eobelidae (two undescribed species), Attelabidae (one
species) and Curculionidae (two species). The first of these is characteristic
of the Mesozoic and is known from the Triassic and Jurassic; the other two
first appeared in the Neocomian. They [Attelabidae and Curculionidae] are
represented by special archaic genera; the two species of Curculionidae
belong to two widely separated branches of the family. Both these branches
occupy a rather special taxonomic position and have already been differen-
tiated by various authors into independent families on the basis of compara-
tive morphological and anatomical studies of the extant forms (Apionidae
and Rhynchophoridae). While the independence of the family Apionidae is
supported by many authors, the separation of the Rhynchophoridae—estab-
lished by Morimoto (1962)—has not yet been widely accepted. In Baisa, the
first group [“Apionidae”’] is represented by the genus Cretonanophyes. The
second apparently includes the genus Slonik, although a strict justification
for its inclusion in the “Rhynchophoridae” is not possible on the basis of the
available material since many important distinguishing characters (structure

182

249

of mouth parts, abdominal terga, genitalia) are not available for study in the
holotype of Slonik sibiricus. If Slonik is in fact included in this group, then
the main line of “"Rhynchophoridae" could have separated from Eobelidae
independent of the other Curculionidae as early as the Late Jurassic or Early
Cretaceous, in which case it definitely merits isolation into a separate family.
On the other hand, “Apionidae” is differentiated from other higher Cur-
culinoidea by several plesiomorphic characters and by only a few secondary
apomorphic ones such as elongation of the trochanters. Hence it is difficult
to be certain of the independence of apionids. Instead they may simply be
regarded as one of the least evolutionarily advanced groups of Curculionidae,
and probably even the ancestors of this family. The discovery that
Cretonanophyes has a rather complete set of plesiomorphic characters for
Curculionidae confirms this interpretation of “Apionidae”. Since the present
material does not provide a basis for dividing Curculionidae into several
families, it seems advisable to tentatively adopt a broad range for this group,
encompassing the “Rhynchophoridae” and “Apionidae”. Therefore,
Cretonanophyes and Slonik are included in Curculionidae. It is hoped that an
investigation of material from other Mesozoic sites will resolve this tangle.

Very little can be said about the ecology of the described species. The
mode of life of Car is unknown although Voss (1953) suggests that it was
associated with conifers; possibly Baissorhynchus developed on conifers as
well. It is impossible to judge the trophic relations of Slonik. There are
grounds for suggesting (in analogy with many extant representatives of
rhynchophoroid stem groups, and based on their general appearance that it
was associated with wood; probably either dead or decomposing wood.
However, in the first place, the inclusion of Slonik in this main line is not
fully substantiated; and secondly, even if it belongs, there are such marked
differences from the extant representatives that an analogy with them might
prove incorrect. With regard to Cretonanophyes, its larvae probably lived on
the generative organs of some angiosperms. Among the present-day
Nanophyinae there is one genus (Nanodiscus Kiesw.) associated with
Juniperus and Thuja, but this relationship is probably secondary. Gall-form-
ing Nanophyinae are known, but they largely belong to the advanced groups
of the subfamily, suggesting that their infestation of fruit is primary. The
spectrum of food plants of representatives of the subfamily is extremely wide
(Ericaceae, Tamaricaceae, Dipterocarpaceae, Lythraceae, Crassulaceae,
Ebenaceae, and others). Therefore, it is impossible to be precise about the
taxonomic position of the host plant of Cretananophyes. Representatives of
one group of insects associated with Angiospermae, viz. Cephidae
(Hymenoptera), have already been described from Baisa (Rasnitsyn, 1969).
Rasnitsyn suggested the existence of angiosperm flora in Baisa ata time when
their remains were still unknown from this site. In 1969, during an expedition
of the Paleontological Institute, Academy of Sciences, USSR, to Baisa,

250

remains of dicotyledons of uncertain taxonomic position were collected
(Vakhrameev, 1973). This discovery was proof of the existence of flowering
plants in the Neocomian of Trans-Baikal, and confirmed the validity of
predictions based on studies of the entomofauna. Insects associated with
angiosperms, and insect-host plant associations in general, are few in Baisa.
The host plants probably occurred there in limited quantities. The activity of
insects favors their burial over that of their host plants, and from the
composition [of the insect fossils] one can quite frequently judge the flora of
territories somewhat removed from the water bodies in which the insects
were buried. In several instances, the composition of the entomofauna may
be the sole source of this type of information.

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PLATES

199

258

PLATE I

Families Triaplidae, Parahygrobiidae and Coptoclavidae

1 and 2. Triaplus macroplatus, sp. nov. (Cc).
1—Holotype PIN No. 2971/104 (x 7.5); 2—Paratype PIN No. 2905/24 —
(x 8); Madygen, Triassic.
3. Triaplus laticoxa, sp. nov. (Cc).
Paratype PIN No. 2555/1720 (x 4.5); Dzhailyaucho, Triassic.
4. Parahygrobia natans sp. nov. (Cc).
Holotype PIN No. 3053/423; a—general view (x 8.0), b—head
(x 20.0); Uda, Jurassic. ©
5. Necronectes aquaticus sp. nov. (Cc).
Holotype PIN No. 722/23; a—general view (x 2.0), b—fore and middle
legs (x 5.0); Ust’-Balei, Jurassic.
6. Necronectes gigas sp. nov. (Cc).
Holotype PIN No. 2554/477 (x 1.1); Karatau, Upper Jurassic.
7. Necronectes cyrenaicus sp. nov. (Cc).
Holotype PIN No. 3394/24 (x 1.4); Bryanka, Lower Cretaceous.
8. ? Necronectes latus sp. nov. (Cc).
Holotype PIN No. 3072/151* (x 5.5); Shurab III, Jurassic.

*Given as 3073/15] in the text and Figure 7—General Editor.

260

PLATE I

Family Coptoclavidae S

1. Exedia plana sp. nov. (c)
Holotype PIN No. 2554/778 (x 1.7); Karatau, Upper Jurassic.

2 and 3. Stygeonectes jurassicus sp. nov. (C)
Paratype: 2—PIN No. 3000/939 (x 10.0); 3—PIN No. 3000/986; a—
(x 6.0), b— Head (x 20.0); Novospasskoe, Jurassic.

4 and 5. Charonoscapha grossa sp. nov. (Cc)
4—Holotype PIN No. 2554/446; a—(x 2.5), b— head and pronotum
(x 4.0); 5—Paratype PIN No. 2997/1847 (x 2.5); Karatau, Upper Juras-
SIC.

6 and 7. Charonoscapha ovata sp. nov. (Cc)
6—Holotype PIN No. 2997/1845 (x 4.0); 7—Paratype PIN No.
2066/2313* (x 3.1); Karatau, Upper Jurassic.

8. Charonoscaphidia elongata sp. nov. (Cc)
Holotype PIN No. 2384/717 (x 2.7); Karatau, Upper Jurassic.

* Given as 2066/2393 in the text and Figure 11—General Editor.

See eee
emer

oe
hae ae
eae

200

262

PLATE Ill

Families Liadytidae, Dytiscidae and Gyrinidae

1. Liadytes longus sp. nov. (c)
Holotype PIN No. 2046/1 (x 2.0); Daya, Lower Cretaceous.
2. Liadytes crassus sp. nov. (Cc)
Holotype PIN No. 3015/369 (x 10.0); Unda, Lower Cretaceous.
3. Cretodytes latipes sp. nov. (c)
Holotype PIN No. 2383/256 (x 6.0); Kyzyl-Dzhar, Upper Cretaceous.
4. Avitortor primitivus sp. nov. (Cc)
Holotype PIN No. 3064/856 (x 6.3); Baisa, Lower Cretaceous.
5 and 6. Mesodyneutes amurensis sp. nov. (Cc)
5—Holotype PIN No. 2055/74 (x 5.0). 6— Paratype PIN No. 2055/79
(x 6.5); Arkhara, Upper Cretaceous.
7. Angarogyrus minimus sp. nov. (Cc)
Holotype PIN No. 1874/30 (x 11.8); Iya, Jurassic.
8. Cretotortor archarensis sp. nov. (Cc)
Holotype PIN No. 2055/75 (x 6.5); Arkhara. Upper Cretaceous.
9. Triadogyrus sternalis sp. nov. (Cc)
Holotype PIN No. 3320/131* (x 5.4); Garazhovka, Upper Triassic.

*Given as 3320/13 in the text and Figure 53—General Editor.

264

PLATE IV

Family Trachypacheidae

1. Sogdodromeus altus sp. nov. (c)
Holotype PIN No. 2971/1417* (x 6.5); Madygen Triassic.
2. Platycoxa armata sp. nov. (c)
Holotype PIN No. 2903/222 (x 7.5); Issyk-Kul’, Lower Jurassic.
3. Platycoxa jurassica sp. nov. (Cc)
Holotype PIN No. 2997/6** (x 9.2); Kenderlyk, Lower Jurassic.
4. Unda microplata sp. nov. (c)
Holotype PIN No. 3015/371 (x 10.0); Unda, Lower Cretaceous.
5. Unda angulata sp. nov. (c)
Holotype PIN No. 2372/24 (x 10.0); a—prothorax; b—metathorax and
abdomen; Daya, Lower Cretaceous.
6. Unda cursoria sp. nov. (Cc)
Holotype PIN No. 2372/23 (x 6.5); Daya Lower Cretaceous.
7. Psacodromeus gutta sp. nov. (c)
Holotype PIN No. 2554/456 (x 5.0); Karatau, Upper Jurassic.
8. Psacodromeus ovalis sp. nov. (c)
Holotype PIN No. 2997/1849 (x 5.7); Karatau, Upper Jurassic.

*Given as 2971/417 in the text and Figure 21—General Editor.
**Given as 2496/6 in the text and Figure 22>—General Editor.

266

PLATE V

Family Trachypacheidae

1. Psacodromeus crassus sp. nov. (Cc)
Holotype PIN No. 2066/3147 (x 8.6); Karatau, Upper Jurassic.
2. Psacodromeus rugosus sp. nov. (c)
Holotype PIN No. 1789/214 (x 4.4); Karatau, Upper Jurassic.
3. Karatoma agilis sp. nov. (c)
Holotype PIN No. 2784/1528 (x 2.5); Karatau, Upper Jurassic.
4. Karatoma raptor sp. nov. (Cc)
Holotype PIN No. 3015/362 (x 5.2); a—reverse impression; b—direct
impression; Unda, Lower Cretaceous.
5. Karadromeus rostratus sp. nov. (c)
Holotype PIN No. 2904/873 (x 8.0); Karatau, Upper Jurassic.
6. Karadromeus latus sp. nov. (Cc)
Holotype PIN No. 2384/567 (x 6.7); Karatau, Upper Jurassic.
7. Karadromeus elongatus sp. nov. (c)
Holotype PIN No. 3015/303* (x 13.7); Unda, Lower Cretaceous.
8. Karadromeus mongolicus sp. nov. (c)
Holotype PIN No. 3145/753 (x 7.8); Anda-Khuduk, Lower Cretaceous.

* Given as 3015/363 in the text and Figure 34a—General Editor.

268

PLATE VI

Families Trachypacheidae and Carabidae

1. Eodromeus antiquus sp. nov. (Cc)
Holotype PIN No. 2239/897 (x 6.5): a—reverse impression; b—direct
impression; Karatau, Upper Jurassic.

201 2. Eodromeus sternalis sp. nov. (c) :

Holotype PIN No. 3191/5 (x 8.0); Chikoiskaya basin, Lower
Cretaceous.

3 and 4. Eodromeus dissectus sp. nov. (Cc)
3—Holotype PIN No. 1989/2976* (x 15.0); 4—Paratype PIN No.
3064/803** (x 7.0); Baisa, Lower Cretaceous.

5. Eodromeus major sp. nov. (c)
Holotype PIN No. 3064/864 (x 4.7); Baisa, Lower Cretaceous.

6 and 7. Protorabus planus sp. nov. (c)
6—Holotype PIN No. 2066/3185 (x 4.1); 7— Paratype PIN No.
2904/928 (x 3.3); Karatau, Upper Jurassic.

8. Protorabus nigrimonticola sp. nov. (Cc)
Holotype PIN No. 2997/1851 (x 5.8); Karatau, Upper Jurassic.

9. Protorabus magnus sp. nov. (Cc)
Holotype PIN No. 2554/ 447 (x 3.0); Karatau, Upper Jurassic.

* Given as 1989/2967 in the text and Figure 37a—General Editor.
** Given as 3064/863 in the text —General Editor.

270

PLATE Vil

Family Carabidae

1. Ovrabites ovalis sp. nov. (c)

Holotype PIN No. 2904/872 (x 4.0); Karatau, Upper Jurassic.
2. Overabites jurassicus sp. nov. (Cc)

Holotype PIN No. 2904/877 (x 10.0); Karatau, Upper Jurassic.
3. Cordorabus notatus sp. nov. (Cc)

Holotype PIN No. 2066/2739 (x 5.0); Karatau, Upper Jurassic.
4. Cordorabus antennatus sp. nov. (Cc)

Holotype PIN No. 2066/2363 (x 4.7); Karatau, Upper Jurassic.
5. Cordorabus minimus sp. nov. (c)

Holotype PIN No. 2239/905 (x 10.0); Karatau, Upper Jurassic.
6. Lithorabus incertus sp. nov. (Cc)

Holotype PIN No. 371/49 (x 16.1); Issyk-Kul’, Lower Jurassic.
7. Cretorabus capitatus sp. nov. (Cc)

Holotype PIN No. 3064/862 (x 9.3); Baisa, Lower Cretaceous.
8. Cretorabus latus sp. nov. (Cc)

Holotype PIN No. 3064/852 (x 3.2); Baisa, Lower, Cretaceous.

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PLATE VII

Family, Carabidae, Adephaga Incertae Sedis

. Mesorabus elongatus sp. nov. (c)

Holotype PIN No. 2784/1525 (x 2.5): a—reverse impression; b—direct
impression; Karatau, Upper Jurassic.

. Conjunctia prodroma sp. nov. (c)

Holotype PIN No. 1668/1763 (x 10.0); Baisa, Lower Cretaceous.

. “Carabites” vitimensis sp. nov. (Cc)

Holotype PIN No. 3064/871 (x 12.0); Baisa, Lower Cretaceous

. “Carabites” creta sp. nov. (Cc)

Holotype PIN No. 2383/111 (x 10.8): a—head and pronotum; b—
metathorax, Kyzyl-Dzhar, Upper Cretaceous.
Harpalinae gen. sp. (c)
specimen PIN No. 2383/206a (x 14.0); Kyzyl-Dzhar, Upper Cretaceous.
Necronectulus avus sp. nov. (Cc)
Holotype PIN No. 1362/27 (x 12.0); Kenderlyk, Lower Cretaceous.

7—9. Cretotaenia pallida sp. nov. (c)

7—Holotype PIN No. 1989/2890: a—general view (x 7.0); b— head
(x 40.0). Paratypes: 8—PIN No. 1989/2889 (x 5.5); 9—PIN No.
1668/1835* (x 5.5); Baisa, Lower Cretaceous.

*Given as 1668/1837 in Figure 52e—General Editor.

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274

PLATE IX

Familes Hydraenidae and Hydrophilidae

1. Peltosyne triassica sp. nov. (Cc)
Holotype PIN No. 2240/278 (x 6.3); Dzhailyaucho, Triassic.
2. “Ochthebiites” altus sp. nov. (c)
Holotype PIN No. 3000/917 (x 20.0); Novospasskoe, Jurassic.
3 and 4. Mesospercheus tarsalis sp. nov. (Cc)
3—Holotype PIN No. 3015/367 (x 8.5); 4—Paratype PIN No. 3063/116
(x 12.5); Unda, Lower Cretaceous.
5. Mesospercheus notatus sp. nov. (Cc)
Holotype PIN No. 3000/923 (x 6.4); Novospasskoe, Jurassic.
6. Mesydra elongata sp. nov. (c)
Holotype PIN No. 1989/2983 (x 10.0); Baisa, Lower Cretaceous.
7. Mesohelophorus sibiricus sp. nov. (Cc)
Holotype PIN No. 3064/841 (x 17.0): a—reverse impression; b—direct
impression; Baisa, Lower Cretaceous.
8. Paraspercheus asiaticus sp. nov. (Cc)
Holotype PIN No. 2997/522* (x 4.7); Karatau, Upper Jurassic.
9. Paraspercheus vitimensis sp. nov. (C)
Holotype PIN No. 3064/1071 (x 4.2); Baisa, Lower Cretaceous.

* Given as 2997/552 in the text —General Editor.

276

PLATE X

Families Silphidae and Scarabaeidae

1-3. Mesagyrtes communis sp. nov. (c)
1—Holotype PIN No. 3000/926 (x 11.0). Paratypes: 2—PIN No.

3000/913 (x 13.0); 3—PIN No. 3000/915 (x 18.0). Novospasskoe,
Jurassic.

4 and 5. Geotrupoides sulcatus sp. nov. (c)
4— Holotype PIN No. 1668/1785 (x 10.0); a—reverse impression; b—

direct impression; 5—Paratype PIN No. 3064/867 (x 10.0); Baisa.
Lower Cretaceous.

6. Geotrupoides leptoscelis sp. nov. (c)
Holotype PIN No. 3064/936 (x 10.0); a—reverse impression; b—direct
impression; Baisa, Lower Cretaceous.

7. Geotrupoides vitimensis sp. nov. (C)

Holotype PIN No. 1668/1805 (x 6.8); Baisa, Lower Cretaceous.

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278

PLATE XI

Family Scarabaeidae

1-4. Proteroscarabeus baissensis sp. nov. (Cc)
1— Holotype PIN No. 1668/1830 (x 1.9). Paratypes: 2—PIN No.
1668/2885* (x 3.0); 3—PIN No. 1668/761** (x 3.0); 4——PIN No.
3064/1072 (x 2.2); Baisa, Lower Cretaceous.

5 and 6. Proteroscarabeus yeni Grabau (c)
5—Specimen PIN No. 2385/2 (x 2.0); Semen depression, Lower
Cretaceous; 6—specimen PIN No. 1989/2638 (x 10.0); Baisa, Lower
Cretaceous.

7. Holcorobeus vittatus sp. nov. (c)
Holotype PIN No. 3064/939 (x 5.4); Baisa, Lower Cretaceous.

8. Holcorobeus picturatus (c)
Holotype PIN No. 1989/2994 (x 4.1); Baisa, Lower Cretaceous.

*Given as 1989/2885 in the text and Figure 68b—General Editor.
** Given as 1668/1761 in the text and Figure 68c—General Editor.

280

203h5*

PLATE XI

Families Cerophytidae, Acanthocnemidae, Cryptophagidae and
Eobelidae

. Aphytocerus communis sp. nov. (C)

Holotype PIN No. 3311/41; a—anterior part of body (x 30.0); b—
genitalia (x 47.0); Yantardakh, Upper Cretaceous.

. Acanthocnemoides sukatshevae sp. nov. (Cc)

Holotype PIN No. 3308/1 (x 27.9); Zhdanikha, Cretaceous.

. Naganasania khetica sp. nov. (c)

Holotype PIN No. 3311/45 (x 28.0); Yantardakh, Upper Cretaceous.

. Succinimontia infleta sp. nov. (Cc)

Holotype PIN No. 3311/31* (45.5): a—ventral view; b—dorsal view;
Yantardakh, Upper Cretaceous.

Eobelus longipes sp. nov. (Cc)
Holotype PIN No. 2452/275 (x 5.4); Karatau, Upper Jurassic.

. Archaeorrhynchus paradoxopus sp. nov. (c)

Holotype PIN No. 2554/715** (x 3.8); Karatau, Upper Jurassic.

* Given as 3130/31 in the text —General Editor.
** Given as 2335/42 in the text and Figure 79—General Editor.

282

6.

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PLATE XII

Family Eobelidae

. Probelus curvispinus sp. nov. (Cc)

Holotype PIN No. 2554/709 (x 6.0).

. Probelopsis acutiapex sp. nov. (Cc)

Holotype PIN No. 2239/1554 (x 5.5).

. Belonotaris punctatissimus sp. nov. (Cc)

Holotype PIN No. 2452/422* (x 2.4).

. Procurculio fortipes sp. nov. (C)

Holotype PIN No. 2066/2339 (x 5.6).

. Eccoptothorax latipennis sp. nov. (c)

Holotype PIN No. 2554/720 (x 6.6)
Oxycorynoides similis sp. nov. (Cc)

Holotype PIN No. 2554/713 (x 15.0)
Scelocamptus tenuirostris sp. nov. (C)

Holotype PIN No. 2554/732 (x 4.5).

All the specimens shown on this plate are from Karatau, Upper Jurassic.

* Given as 2462/422 in Figure 85—Gencral Editor.

284

PLATE XIV

Families Eobelidae, Attelabidae and Curculionidae

1. Ampliceps dentitibia sp: nov. (c)

Holotype PIN No. 2239/1562 (x 17.2); Karatau, Upper Jurassic.
2. Paroxycorynoides elegans sp. nov. (c)

Holotype PIN No. 2452/125 (x 10.0); Karatau, Upper Jurassic.
3. Eccoptarthrus crassipes sp. nov. (c)

Holotype PIN No. 2239/1507 (x 7.5); Karatau, Upper Jurassic.
4. Distenorrhinus angulatus sp. nov. (c)

Holotype PIN No. 2239/1547 (x 12); Karatau, Upper Jurassic.
5. Brenthorrhinus mirabilis sp. nov. (c)

Holotype PIN No. 2554/708 (x 7.0); Karatau, Upper Jurassic.
6. Nanophydes ovatus sp. nov. (c)

Holotype PIN No. 1789/102 (x 14.3); Karatau, Upper Jurassic.
7. Baissorhynchus tarsalis sp. nov. (c)

Holotype PIN No. 1989/3010 (x 18.2); Baisa, Lower Cretaceous.
8. Cretonanophyes longirostris sp. nov. (c)

Holotype PIN No. 1668/1772 (x 12.3); Baisa, Lower Cretaceous.
9. Slonik sibiricus sp. nov. (c)

Holotype PIN No. 1989/2938 (x 17.7); Baisa, Lower Cretaceous.

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