FIRST RECORD OF A FOSSIL BEETLE
(COLEOPTERA, HALIPLIDAE) FROM THE BASAL PALEOCENE
FLYSCH SEDIMENTS IN THE MAGURA UNIT (OUTER WESTERN
, ANDRÉ NEL
, JIØÍ HÁJEK
and MIROSLAV BUBÍK
Department of Zoology, Charles University, Vinièná 7, 128 44 Praha 2, Czech Republic; firstname.lastname@example.org
Laboratoire dEntomologie and CNRS UMR 5143, Muséum National dHistoire Naturelle, rue Buffon 45, F-750 05 Paris, France;
Czech Geological Survey, Leitnerova 22, 602 00 Brno, Czech Republic
(Manuscript received May 28, 2003; accepted in revised form December 16, 2003)
Abstract: The first record of an insect (Coleoptera: Adephaga) from the base of Paleocene marine deposits in the Outer
Flysch Carpathians of eastern Moravia (Czech Republic) is described. Its familial attribution to Gyrinidae or Haliplidae
is critically discussed in comparison to fossil and recent representatives. The presence of a beetle elytron from hemipelagite
in Early Paleocene deep-sea deposits is significant for taphonomy and suggests long wind transport from lagoonal to
open marine environment and disarticulation by predators or scavengers in surface waters.
Key words: Paleocene, Outer Western Carpathians, taphonomy, Insecta, Coleoptera, Haliplidae, Gyrinidae.
Archaic forms of Adephaga were possibly present in the Late
Paleozoic beetle faunas, but this group is accurately known
from the Mesozoic. Extinct Triaplidae or recent Trachy-
pachidae were already present during the Triassic time (Pono-
marenko 1977). During the Jurassic period, the number of
Adephaga rapidly increased up to 10 % of the total diversity
of Coleoptera, represented by the families Carabidae, Colym-
bothetidae, Coptoclavidae, Dytiscidae, Gyrinidae, Jurodidae,
Liadytidae, Parahygrobiidae, Trachypachidae and Triaplidae
(Ponomarenko 1995). The Early Cretaceous assemblages are
similar to those from the Jurassic deposits with diverse Trachy-
pachidae, while the Late Cretaceous beetle faunas are closer to
those from the Cenozoic (Ponomarenko 1995).
The Gyrinidae and Haliplidae are regarded as the two most
basal lineages of Adephaga (or Hydradephaga) (Beutel 1995),
which implies a great antiquity for these families.
Fossil whirligig beetles (Gyrinidae) are known from the Up-
per Triassic until the present and they are the most common
water beetles during the Late Cretaceous (Ponomarenko 1977,
1995). The Mesozoic diversity of Gyrinidae is higher than that
of the Dytiscidae, which is the dominant group of aquatic bee-
tles in many Cenozoic deposits. Another feature of the Meso-
zoic aquatic beetle fauna is the more frequent presence of
brackish and lagoonal inhabitants with respect to the recent fau-
na. Correlation between continental and marine deposits using
Mesozoic beetles is a possible future perspective (Ponomarenko
et al. 1999). A sparse record of fossil crawling water beetles
(Haliplidae) is first known from Lower to middle Cretaceous
deposits (Dmitriev & Zherikhin 1988; Ren et al. 1995).
The further record of Haliplidae is still very poor. The genus
Haliplus is recorded from the Miocene of Oeningen (Germa-
ny) (Schöberlin 1888; Guignot 19311933); Ponomarenko
(1969) listed an isolated elytron from the late Lower Creta-
ceous of Labrador (Canada) that bears a strong resemblance to
the haliplid Peltodytes, Ren et al. (1999) listed the family
Haliplidae from the Late Mesozoic of Hebei Province, China;
and Ren et al. (1995) described Cretihaliplus chifengensis and
Cretihaliplus sidaojingensis from the Early Cretaceous of In-
ner Mongolia, China.
The single find of an insect was made during field research
of the K/T boundary section near Uzgruò by one of the au-
thors (M. Bubík). The Uzgruò K/T section is situated on the
left bank of an unnamed creek NNE of the Uzgruò settlement
near Velké Karlovice close to the Czech-Slovak border
(Fig. 1). The sediments in the Uzgruò vicinity are part of the
Raèa Unit of the Magura Group of Nappes, Outer Flysch Car-
pathians. The insect bearing strata are assigned to the Soláò
The sedimentary succession across the K/T boundary near
Uzgruò can be characterized as fine alternation of muddy tur-
bidites and hemipelagites. Few thicker sandy turbidites (up to
40 cm thick) with missing fine part occur close to the K/T
boundary. The majority of sandstones are up to a few cm thick
and laterally they disappear over a short distance (Tc ripple
sandstones). The most common turbidites comprise Te, Tde,
and less frequently Tcde intervals of the Bouma classification.
Across the K/T boundary a coarsening-upward trend was ob-
served in both small and large scale (Bubík et al. 1999). The
prevailing lithologies are grey-green, grey, and dark grey clay-
stones intercalated by siltstones and fine- to medium-grained
GEOLOGICA CARPATHICA, 55, 6, BRATISLAVA, DECEMBER 2004
470 PROKOP, NEL, HÁJEK and BUBÍK
Fig. 1. Geographical and tectonic situation of the Uzgruò section
(after Bubík et al. 1999, modified). A Tectonic map of the Out-
er Flysch Carpathians. The situation of the detail map B is marked
by arrow. B Geological map of the area around the Uzgruò sec-
tion; situations of sections 19 and 20 marked by arrows.
graywacke sandstones. Less frequently, thin banks of marl-
stones and rarely pelocarbonates occur. Marlstones contain up
to 45 % of carbonate. Turbidite claystones usually have less
than 10 % of carbonate. Hemipelagic claystones are solely
non-calcareous. Carbonates and calcareous claystones are re-
stricted to the Maastrichtian part of the section. Paleocene sed-
iments of the Soláò Formation in the Uzgruò area are strictly
The K/T boundary multistratigraphy of the section was re-
cently established by Bubík et al. (2002) using geochemical
anomalies and all available microfossil groups (foraminifers,
radiolarians, calcareous nannofossils, and dinocysts). The K/T
boundary layer itself, known as a worldwide distributed cor-
relative horizon recognized by the Ir anomaly, shock quartz,
and other impact-related features, was not found at the Uzgruò
section. The layer was probably removed by submarine ero-
sion. However, the weak increase in Ir concentrations was de-
tected in three closely spaced muddy turbidites. This is inter-
preted as redeposition of the Ir-bearing boundary layer. The K/T
boundary in the section is then tentatively defined at the base
of the lowest Ir-bearing turbidite. This is in agreement with the
FOSSIL BEETLE (COLEOPTERA) FROM PALEOCENE FLYSCH SEDIMENTS (MORAVIA) 471
Fig. 2. Haliplidae gen. et sp. indet., Paleocene, Soláò Formation,
Uzgruò, right elytron ventral view (T02729, National Museum
Prague coll.). A photograph. B line-drawing.
biostratigraphic data. The last occurrence of Maastrichtian
nannofossils and planktonic foraminifer Abathomphalus maya-
roensis was recorded 65 cm below this level and the Pale-
ocene dinocyst index Carpatella cornuta (Dinophyceae) ap-
pears just 35 cm above.
The beetle elytron was found in green-gray hemipelagite
claystone 25 to 30 cm above the K/T boundary in the basal Pa-
leocene. Diversified flysch-type agglutinated foraminiferal
fauna, dinocysts dominated by oceanic forms, and radiolarians
in this sediment characterize the paleoenvironment as deep-
sea open marine and persistently below the CCD. Besides the
turbidite currents, the bottom was disturbed by bottom cur-
rents as indicated by silty intercalations without sharp base
and with negative gradation. The paleoenvironment was fur-
thermore influenced by an oxygen minimum zone in the water
column and possible weak upwelling (Bubík et al. 2002). The
oxygenated bottom conditions persisted across the K/T
boundary as shown by the benthic foraminiferal fauna and pa-
leoichnological record (Bubík et al. 2002).
Suborder: Adephaga Emery, 1886
Family: Haliplidae Thomson, 1860
Haliplidae gen. et sp. indet.
Description: Elytron 4.9 mm long, 2.6 mm wide, ratio
length to greatest width 1.88, ratio of length to basal elytral
width 3.1; six distinct rows of large irregular punctures, each
row with maximally 17 punctures; no visible polygonal pat-
tern but micropunctures present on entire surface; sutural stria
not present and elytral suture not deflected near apex; lateral
edge of elytron weakly sinuate distally; epipleuron not recog-
Material: Specimen T02729 (National Museum Prague
coll.), imprint and counter-imprint of elytron with well-pre-
served structure and visible six puncture rows in dorsal view.
Age and layer: Paleocene, Soláò Formation, Uzgruò,
Discussion: The present specimen has some characteris-
tics of both the families Gyrinidae and Haliplidae in the struc-
ture and shape of the elytron. These are the presence of six dis-
tinct rows of irregular punctures; lateral edge of elytron
weakly sinuate distally and ratio (3.1) of elytral length to the
basal elytral width sensu Lawrence & Newton (1995). It is ex-
tremely difficult to determine the family affinities of a Co-
leoptera on the sole basis of the elytral structures. But, after
Khalav (1980), the Haliplidae have no polygonal pattern on
the surface of the elytron, unlike the Gyrinidae. This character
suggests that our fossil belongs to the Haliplidae rather than
the Gyrinidae, which is also supported by the elytron size.
We are not able to attribute this fossil to a recent genus be-
cause of the lack of the characters currently used in the taxon-
omy and phylogeny of the family (Chandler 1943; Beutel &
Ruhnau 1990). Nevertheless, our fossil has an ornamentation
of the elytra similar to Peltodytes Régimbart, 1878, that is a
pronounced outer anterior angle and with 67 rows of punc-
tures. But this similarity could be due to a convergence, and it
is not sufficient to attribute this Paleocene fossil to the recent
The fossil Cretihaliplus chifengensis has strong continuous
striae on the upper surface of its elytra, unlike our fossil.
However, there is no fossil or recent Gyrinidae with a pat-
tern of irregular rows of large punctures on elytra. Also, nearly
all the recent Gyrinidae have elytra more or less posteriorly
truncate, but not all of them (Dineutes politus Mac Leay,
1825, see Régimbart 1902: Fig. 1). Therefore affinities with
the Gyrinidae can reasonably be excluded, but a comparison
with the described species is added.
Eleven fossil genera of Gyrinidae have been described until
now. Seven of them are known from the Mesozoic (Ponomar-
enko 1985; Nel 1989; Carpenter 1992). The Lower Jurassic
monotypic Anagyrinus Handlirsch, 1906, with A. atavus
(Heer, 1865), is a little-known genus from the European de-
posits (Germany and Switzerland). Its elytra are without
sculptures except for a marginal line that is different from our
specimen. The Middle Jurassic monotypic Angarogyrus Pono-
marenko, 1977, with A. minimus Ponomarenko, 1977 from Iya
(Cis-Baikal, Irkutsk area in Asian Russia), differs from our
specimen in its smooth and apically rounded elytra. The Low-
er Cretaceous monotypic genus Avitortor Ponomarenko,
1977, with A. primitivus Ponomarenko, 1977 from Baissa
(Transbaikalia, Russia), differs in the presence of fine grooves
unlike the irregular puncture rows of our specimen. The other
Lower Cretaceous monotypic Baissogyrus Ponomarenko,
1973, with B. savilovi Ponomarenko, 1973, is known from
eastern Russia. It is based on a ventral part of the body. Thus,
it is not possible to compare with our specimen. The type spe-
cies C. zherichini Ponomarenko, 1973 of the Upper Creta-
472 PROKOP, NEL, HÁJEK and BUBÍK
ceous genus Cretotortor Ponomarenko, 1973 (Kzyl-dzhar
Karatau, southern Kazakhstan), based on an elytron with 9
prominent furrows and a sutural margin, is different from our
specimen. The same genus comprises C. archarensis Pono-
marenko, 1977 from Arkhara in Russia. Its elytron differs
from our specimen in the sutural margin, distinct transversal
rugae and relatively dense punctuation. From the same Late
Cretaceous locality Arkhara also comes the monotypic Me-
sodineutes Ponomarenko, 1977, with M. amurensis Pono-
marenko, 1977, characterized by almost smooth elytra with
dense ventral punctuation, and distinct margin along the su-
ture. The Mesozoic Mesogyrus Ponomarenko, 1973, with M.
antiquus Ponomarenko, 1973 known from the Late Jurassic of
Kazakhstan, has 89 distinct furrows and a sutural margin.
Other Cretaceous species of this genus are M. sibiricus Pono-
marenko, 1985 and M. striatus Ponomarenko, 1973 from the
Asian part of Russia, which have similar structure of elytra
There are several Tertiary gyrinid genera described (see list
in Nel 1989). Gyrionides Motschulsky, 1856, with G. limba-
tus Motschulsky, 1856 from the Early Oligocene Baltic am-
ber, is similar to the recent genus Gyrinus Linnaeus, 1767,
differing mainly in the absence of striae. It is the smallest
known fossil Gyrinidae (Hatch 1927). Miodineutes Hatch,
1927, with M. oeningensis Hatch, 1927 from the Upper Mi-
ocene of Oeningen (Baden, Germany), is similar to the recent
Enhydrus Laporte de Castelnau, 1834. The elytra of this spe-
cies are without sculpture except for the marginal line. Orec-
tochilus sp. from the Paleocene of Menat (Puy-de-Dôme,
France), lacks similar elytra punctuation. Fourth, Gyrinus
aquisextanea Nel, 1989, described from the Upper Oligocene
of Aix-en-Provence (France), is attributed to a recent genus.
This species has no distinct punctuation, unlike our elytron.
Finally, one Quaternary monotypic Protogyrinus Hatch,
1927, with P. confines (Scudder, 1900) from the Pleistocene
of Ontario (Canada), is also similar to the recent Gyrinus but
differ by presence of rows of punctures and elytral striae ex-
tending independently to the apex (Hatch 1927).
In spite of the fact that we have only a single elytron with-
out strict autapomorphies with Gyrinidae or Haliplidae, we
propose to attribute this specimen to the Haliplidae because of
the unique character of six irregular rows of large punctures
similarly present in recent genus Peltodytes Régimbart, 1878
(67 rows of punctures). This character is not present in any
fossil or recent species of Gyrinidae worldwide. In this fami-
ly, different kinds of punctures are present in superficial stri-
ae, usually much less pronounced than in this fossil and Hali-
plidae. However, any of these characters cannot be used to
make more precise determination or establish a new taxon, so
we propose to leave this specimen in open nomenclature. Fur-
ther findings of a complete body could help to solve the sys-
tematic position of this enigmatic beetle.
Although the systematic position of this elytron is uncer-
tain, the discovery has undoubted taphonomical significance.
It is probably the first insect record from the Magura Flysch of
the Carpathian Orogen at all. This is not surprising when con-
sidering the paleoenvironment of Magura Flysch sediments:
bathyal to abyssal oceanic basin with oxic bottom conditions.
Although terrigenous phytodetritus is a relatively common
component of turbidites of many Cretaceous to Paleogene for-
mations in the Magura Flysch, phytodetrit accumulations
have revealed no insect remains yet. Repeated transport and
deposition in the marginal marine to deep-sea environments is
probably rather harsh processes for preservation of insect ex-
oskeletons. The described beetle elytron was found in hemi-
pelagite sediment deposited in the lower bathyal to abyssal
zone. This rather excludes the transport by turbidite current
and suggests probably long wind transport from lagoonal to
open marine environment and disarticulation by predators or
scavengers in surface waters. The presence of an oxygen min-
imum zone and relatively rapid sedimentation in a turbidite
fan system probably served as favourable conditions for pres-
Acknowledgments: We are grateful to Jozef Michalík (Geo-
logical Institute, Slovak Academy of Sciences), Alexander
Rasnitsyn (Russian Academy of Science), and Peter Vranský
(Geological Institute, Slovak Academy of Sciences) for their
revision and suggestions to the manuscript. The research on
the K/T boundary at Uzgruò was supported by grant of the
Grant Agency of Czech Republic: No 205/00/0218 Pale-
ocene boundary events in the Magura Flysch of Moravia.
The first author acknowledges of the research support to the
Grant Agency of the Czech Republic No. 205/03/D151 and
grant of the Ministry of Schools CEZ: J13/98: 113100004.
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