GEOLOGICA CARPATHICA, APRIL 2007, 58, 2, 181—184

www.geologicacarpathica.sk

Oligaeschna kvaceki sp. nov., a new fossil dragonfly

(Odonata: Aeshnidae) from the middle Oligocene sediments

of northern Moravia (Western Carpathians)

JAKUB PROKOP

1

, TOMÁŠ PŘIKRYL

2

, ONDŘEJ DOSTÁL

3

 and ANDRÉ NEL

4

1

Department of Zoology, Charles University in Prague, Viničná 7, CZ-128 44 Praha 2, Czech Republic;  jprokop@natur.cuni.cz

2

Institute of Geology and Paleontology, Charles University in Prague, Albertov 6, CZ-128 43 Praha 2, Czech Republic;  prikryl.t@seznam.cz

3

Department of Paleontology, Masaryk University, Kotlářská 2, CZ-311 37 Brno, Czech Republic;  dostal.muzeum@boskovice.cz

4

CNRS UMR 5202, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 rue Buffon, F-75005 Paris, France;  anel@mnhn.fr

(Manuscript received March 31, 2006; accepted in revised form October 5, 2006)

Abstract: A new species of fossil dragonfly Oligaeschna Piton et Théobald, 1939 (O. kvaceki sp. nov.) is described
from Middle Oligocene strata of northern Moravia and compared with all closely related species. The current record
documents a rather broad distribution and probably also high abundance of Oligaeschna in Eurasia during the
Oligocene and Miocene.

Key words: Middle Oligocene, Western Carpathians, taxonomy, Insecta, Aeshnidae, Aeshninae.

Introduction

Fossil insects in the territory of Western Carpathians are
mainly known from the Czech Republic and Slovakia by
scarce fauna from Paleogene and Neogene deposits and
amber resin (e.g. Říha 1979; Prokop et al. 2004; Prokop &
Nel 2005; Sukatcheva et al. 2006).

Kalabis (1950) first noticed the fossil insects from the

Kelč locality near Hranice in Moravia. He reported a lost
specimen of fossil bug (Heteroptera: Alydidae), lately de-
scribed as Sulcalydus kalabisi Štys et Říha, 1975. This spe-
cies is based on remarkable photographs only, because the
original specimen has been lost in the Moravian Museum
in Brno during the Second World War (Štys & Říha 1975).

Fossils at the Kelč locality are preserved in yellow-grey

fine sandy claystones with calcareous admixture and
marked lamination of the middle Oligocene (Ruppelian/
Chattian) Menilite Formation (Western Carpathians, late
Tethys Sea). The microbiostratigraphy of samples from the
locality based on calcareous nannoplankton are still in
preparation. However the general data on phytostratigra-
phy could be established on the basis of the published
macroflora (Knobloch 1969). Macrofauna is also well doc-
umented by abundant fish remains primarily studied by
Kalabis (1975a,b) and summarized by Přikryl (2004). Be-
side the sparse insect record, the locality is famous for
abundant fish fossils counting seven genera of teleostean
fishes (Clupea Linnaeus, 1758; Glossanodon Guichenot,
1867;  Scopeloides Wettstein, 1886; Vinciguerria Good et
Bean, 1895; Diaphus Eigenmann et Eigenmann, 1890;
Anenchelum Blainville, 1818; Serranus  Cuvier, 1817) and
two genera of Elasmobranchii (Alopias  Rafinesque, 1810;
Cetorhinus  Blainville, 1816) indicating an epipelagic to
mesopelagic paleoenvironment. The only genus Serranus
could also be classified as a classical representative of a

neritic paleoenvironment (Přikryl 2004). The original ma-
terial studied by Kalabis was collected at the “Kelč-
zámek” (= Kelč-chateau) locality, nowadays inaccessible
due to building. The new material comes from the Kelč-
Strážné locality (Přikryl 2004).

We follow the phylogeny and classification proposed

by Ellenrieder (2002) and wing venation nomenclature of
Riek & Kukalová-Peck (1984) and Kukalová-Peck (1991)
as amended by Nel et al. (1993) and Bechly (1996).

Systematic Paleontology

The used abbreviations see in caption to Fig. 1. Paleon-

tological material is deposited in the Museum of Valašs-
ko Region in Vsetín (Czech Republic).

Family:  Aeshnidae  Leach, 1815

Genus:  Oligaeschna Piton et Théobald, 1939

T y p e   s p e c i e s :

  Oligaeschna  jungi Piton et Théobald,

1939.

Oligaeschna kvaceki sp. nov., Prokop et Nel

(Fig. 1A—B)

D i a g n o s i s :

 Differing from other Oligaeschna  species

in the discoidal triangle rather broad, divided into five
small cells, and as long as supratriangle; arculus aligned
with basal side of discoidal triangle; pterostigma covering
five cross-veins between RA and RP1; RP2 evenly curved
two cells basal of proximal margin of pterostigma; two rows
of cells between MA and Mspl along the posterior wing
margin; CuAb directed towards base of wing and partly pos-
teriorly closing anal loop with AA; anal area rather narrow.

182

PROKOP, PŘIKRYL, DOSTÁL and NEL

D e s c r i p t i o n :

 Hind wing about 50 mm long and

18 mm wide; distance from base to nodus about 21 mm;
estimated distance from nodus to apex about 29 mm; dis-
tance from nodus to pterostigma 18.4 mm; estimated dis-
tance from pterostigma to apex 4.6 mm; pterostigma
rather long, 5.2 mm long and 0.9 mm wide; oblique
pterostigmal brace cross-vein present and aligned with
base of pterostigma; 17 postnodal cross-veins, not well
aligned with corresponding subpostnodal cross-veins; 15
visible secondary antenodal cross-veins not aligned with
corresponding cross-veins between ScP and RA; six sec-
ondary subantenodal cross-veins between Ax2 and nodus
and 5 between Ax2 and Ax1; Ax1 two cells basal of arcu-
lus, 7.4 mm from wing base; Ax2 two cells basal of distal

discoidal angle, 13.9 mm from wing base; numerous cross-
veins between RA and RP basal of subnodus; median
space free; supratriangle narrow elongate, not visibly
crossed by any veins; arculus aligned with basal side of
discoidal triangle; submedian space crossed by four cross-
veins, including CuP; PsA not stronger nor more oblique
than other cross-veins in submedian space; discoidal trian-
gle rather broad and divided into five small cells, its costal
side 5.0 mm long and proximal side 2.8 mm long; RP2
evenly curved two cells basal of proximal margin of
pterostigma; Rspl not parallel to IR2, with a concave bend
in its distal part, and four rows of cells between it and IR2
near bending; MA with a marked concave bend but no
well-defined secondary longitudinal vein in distal part of

Fig. 1.  Oligaeschna kvaceki sp. nov., holotype No. 235/05. A – drawing of female hind wing (Ax1 – basal primary antenodal cross-
vein, A x2 – distal primary antenodal crossvein, ScP – subcosta posterior, RA – radius anterior, RP1 – radius posterior 1, IR1 – in-
terradius 1, RP2 – radius posterior 2, IR2 – interradius 2, Rspl – radial supplement, RP1 – radius posterior 3 + 4, MA – media ante-
rior, Mspl – median supplement, MP – media posterior, CuAa – the main branch of cubitus anterior, CuAb – the first posterior
branch of cubitus anterior, AA1b – the first posterior branch of analis anterior, AA2b – the second posterior branch of analis anterior,
CuP – cubitus posterior, PsA – pseudo-analis). B – photograph of hind wing imprint (scale bar represents 5 mm).

183

OLIGAESCHNA KVACEKI SP. NOV., A NEW OLIGOCENE FOSSIL DRAGONFLY (WESTERN CARPATHIANS)

area between RP3/4 and MA; one or two rows of cells be-
tween MA and RP3/4 in distal part of this area; Mspl well
defined, with a slightly concave bend in its distal part;
three rows of cells between Mspl and MP in postdiscoidal
area and about eleven near hind margin of wing; four rows
of cells between Mspl and MA in widest part; oblique
cross-vein ‘O’ slightly distal (two cells) of base of RP2;
CuAa with more than four posterior branches directed to-
wards hind margin of wing; cubito-anal area 5.5 mm wide;
CuAb directed towards base of wing and partly (only two
cells long) posteriorly closing anal loop with AA; anal
loop rather narrow with three rows of cells, and high with
five or six rows of cells, 4.8 mm high, 2.8 mm wide, and
divided into 10 visible cells (rather incomplete with origi-
nally about 14 cells); anal area narrow, 4.8 mm wide, with
two or three rows of cells and two main veins AA1b and
AA2b, postero-basal margin rounded (female hind wing).

H o l o t y p e :

 Specimen No. 235/05 (almost complete fe-

male hind wing with extreme base and apex missing, well
preserved imprint and counterprint), deposited in the Mu-
seum of Valašsko Region in Vsetín, north-eastern Czech
Republic.

T y p e   l o c a l i t y :

 Kelč-Strážné západ (Moravia), Czech

Republic.

T y p e   s t r a t a :

 Middle Oligocene (Ruppelian/Chatti-

an), “Menilite Formation” (yellow-grey fine sandy clay-
stones with calcareous admixture and marked lamination).

E t y m o l o g y :

 Named in honour of Zlatko Kvaček, pa-

leobotanist from Charles University in Prague.

D i s c u s s i o n :

 This hind wing clearly belongs to Aesh-

nidae and to the clade ‘Aeshninae’ sensu Ellenrieder
(2002: fig. 20) because of combination of the following
apomorphic characters: MA and RP3/4 not parallel, MA
with a marked concave bend; Mspl with a concave bend
in its distal portion (homoplasy); Rspl not parallel to IR2,
with a concave bend.

It also has the apomorphy of the group ‘Aeshninae’ mi-

nus (Polycanthagyna & Basiaeshna & Amphiaeshna), viz.
space between MP and CuA at basal third twice as wide as
at distal third, MP and CuA converging towards wing mar-
gin (Ellenrieder 2002: fig. 20). The group (Heliaeschna  &
Neuraeschna & Staurophlebia) can be excluded because
of the absence of a pterostigmal brace and median space
crossed.  Indaeschna Fraser, 1926 also has its median space
crossed (Fraser 1926). The group (Anaciaeschna & Anax
&  Hemianax) can be excluded because of the absence of a
very strong bend of RP2. Among the other genera, Platty-
cantha Förster, 1900 and Argyrtacantha Lieftinck, 1937
have their fork of IR2 in a very proximal position. Tria-
canthagyna Selys, 1883, Gynacantha Rambur, 1842 and
Subaeschna Martin, 1909 have only one row of cells in
the proximal part of the area between the MP and CuA in
the hind wing. Austrogynacantha Tillyard, 1908 and
Staurophlebia Brauer, 1865 have longer and narrower dis-
coidal triangles. Racenaeschna Calvert, 1958 has 2/3 rows
of cells in area between MA and Mspl (Calvert 1958). All
these genera have their IR2 forked, as for Castoraeschna
Calvert, 1952. The fork of IR2 is weak but present, oppo-
site mid part of pterostigma in Coryphaeschna William-

son, 1903, Remartinia Navas, 1911, and Oreaeschna Lieft-
nick, 1937. The possible affinities with genus Aeshna spp.
can be excluded because all the Aeshna spp. have at least
a weak fork of IR2, with an anterior branch not zigzagged.
Also, the RP2 of our fossil has a weak distal concavity, un-
like the Aeshna spp.

Our fossil is closely related to the two genera Oplonae-

schna Selys, 1883 (with two recent species O. armata
(Hagen, 1861) and O. magna Gonzalez Soriano et Novelo
Gutierrez, 1998 and several doubtful fossils, see below),
and fossil Oligaeschna Piton et Théobald, 1939. These
genera differ in the length of pterostigma, distinctly long-
er in Oligaeschna than Oplonaeschna, and structure of
discoidal triangle, longer and divided into five or more
cells in Oligaeschna (except for O. saurai Peñalver et al.,
1996), instead of four in Oplonaeschna (Gonzalez Soriano
et Novelo Gutierrez, 1998). On the basis of the combina-
tion of the above mentioned characters we attribute our
fossil to Oligaeschna.

Nel et al. (1994) revised the genus Oligaeschna and

considered it as the possible sister genus of Oplonaeschna.
At that time, Oplonaeshna was included in the Gom-
phaeschnini, but recently transferred to the ‘Aeshninae’ by
Ellenrieder (2002). Oligaeschna  has  the main venational
synapomorphy of this clade, viz. MA with a concave
bend, and Mspl and Rspl distinctly concave. That is why
Nel et al. (2005) proposed to transfer this genus to the
‘Aeshninae’.

Among the fossil taxa currently attributed to Oplonae-

schna,  O. staurophlebioides Henriksen, 1922 from Upper
Oligocene or Lower Miocene strata of Denmark is based
on an incomplete hind wing that Nel et al. (1994) consid-
ered to be an Aeshnidae incertae sedis.  O. vectensis Cock-
erell et Andrews, 1916 from the Late Eocene of UK and O.
metis (Heer, 1849) from the Early Miocene of Croatia are
also both regarded as an Aeshnidae incertae sedis.

The  Oligaeschna  genus comprises five described spe-

cies from Oligocene and Miocene Eurasian deposits. O.
jungi  Piton et Théobald, 1939 is originally based on fore
wings and a fragment of hind wing from the Oligocene lo-
calities of Puy-de-Mur, Puy-de-Dôme (France), and also re-
corded from the Oligocene section of Enspel (Germany)
and the Lower Oligocene strata of Provence (France)
(Wedmann 2000; Nel et al. 2005). It differs from O. kvace-
ki sp. nov. by more slender discoidal triangle, longer basal
part of CuAb posteriorly closing the anal loop, and
straighter proximal part of RP2.

Both  O. lapidaria (Cockerell et Counts, 1913) (in Cock-

erell 1913) and O. separata (Scudder, 1890), from Lower
Oligocene Florissant fossil site (Colorado, USA), original-
ly attributed to Oplonaeschna, differ from O. kvaceki sp.
nov. as follows: the presence of only three cross-veins be-
tween RA and RP1 below pterostigma instead of five in O.
kvaceki; the arculus basally shifted one cell basal of basal
side of discoidal triangle, so that the discoidal triangle is
distinctly shorter than the supratriangle instead of being
of about equal size as in O. kvaceki (see Cockerell 1913).

O. palaeocoerulea (Timon-David, 1946) from the Oli-

gocene Camoins-les-Bains locality (Bouches-du-Rhône,

184

PROKOP, PŘIKRYL, DOSTÁL and NEL

France) has circa 10 rows of cells between RP2 and IR2 along
wing margin instead of about seven in O. kvaceki, two rows
of cells between RP3/4 and MA and one row of cells between
MA and Mspl along the posterior wing margin instead of op-
posite situation in O. kvaceki (see Nel et al. 1994).

O. ashutasica (Martynov, 1929) from the Paleocene to

Oligocene strata of Ashutas Mount (East Kazakhstan) is
based on a male hind wing that rather closely resembles
our species. It differs by the arculus basally shifted one
cell basal of basal side of discoidal triangle similarly to O.
lapidaria, so that the discoidal triangle is distinctly short-
er than the supratriangle instead of being of about equal
size as in O. kvaceki; much broader anal area instead of be-
ing rather narrow as in O. kvaceki; MA less bent than in O.
kvaceki (see Martynov 1929: fig. 4).

O. saurai Peñalver, Nel et Martínez-Delclós, 1996, from

the Lower Miocene Ribesalbes locality (Castellón, Spain),
is a distinctly smaller species than all the others. Its discoi-
dal triangle is almost in the shape of an equilateral trian-
gle instead of being elongated as in O. kvaceki (could be
due to secondary deformations); CuAb is longer and cov-
ers three cells instead of two in O. kvaceki; the pterostigma
is distinctly smaller, covering three cross-veins between
RA and RP1.

Conclusion

The current record documents a rather broad distribu-

tion and probably also high abundance of Oligaeschna  in
the Oligocene and Miocene deposits of Eurasia.

The fish assemblages from the Kelč-Strážné locality in-

dicate an epipelagic to mesopelagic environment (Přikryl
2004).

Acknowledgments:

 We are grateful to Miloslav Požár

(Museum of Valašsko Region in Vsetín) who collected
and deposited the type specimen in the collection of Mu-
seum of Valašsko Region in Vsetín. The species name is
dedicated to 70th birthday of Prof. Zlatko Kvaček, leader
of the Czech paleobotany at Charles University. We thank
the anonymous reviewers for valuable comments and cor-
rections on the manuscript. The first author acknowledges
the research support from the Grant Agency of the Czech
Republic (Nos. 205/03/D151, 205/05/0204) and a Grant
from the Ministry of Education MSM 0021620828.

References

Bechly G. 1996: Morphologische Untersuchungen am Flügelgeäder

der rezenten Libellen und deren Stammgruppenvertreter (In-
secta; Pterygota; Odonata), unter besonderer Berücksichtigung
der Phylogenetischen Systematik und des Grundplanes der
*Odonata.  Petalura, Böblingen, Spec. Vol. 2, 1—402.

Calvert P.P. 1958: Resultados zoologicos de la expedicion de la

Universidad Central de Venezuela a la region del Aruyantepui
en la Guyana Venezolana. pabirl de 1956. 4. Genus Racenae-
schna new genus (Odonata: Aeshnidae). Acta Biol. Venezuela-
na 2, 20, 227—234.

Cockerell T.D.A. 1913: Two fossil insects from Florissant with a

discussion of the venation of the aeschnine dragon-flies. Proc.
US Natur. Mus. 45, 577—583.

Gonzalez Soriano E. & Novelo Gutierrez R. 1998: Oplonaeschna

magna sp. nov. (Odonata: Aeshnidae), from Mexico with a
description of its larva. Rev. Biol. Tropical. 46, 3, 705—715.

Ellenrieder N. 2002: A phylogenetic analysis of the extant Aesh-

nidae (Odonata: Anisoptera). Sys. Entomol. 27, 437—467.

Fraser F.C. 1926: Notes on a collection of dragonflies (Odonata)

from the Dutch East Indies and descriptions of four new spe-
cies from the neighbouring continent. Treubia 8, 467—494.

Kalabis V. 1950: New paleontological findings from the Paleogene

in Moravia. Přírodovědný sborník Ostravského kraje 11, 322—
323 (in Czech).

Kalabis V. 1975a: Macropaleontological documentation of Meni-

lite Beds concerning especially the ichtyofauna of  Špičky u
Hranic na Moravě and Kelč localities. Part I: Špičky. Zpr.
Vlastivěd. Úst. v Olomouci 173, 1—11 (in Czech).

Kalabis V. 1975b: Macropaleontological documentation of Meni-

lite Beds concerning especially the ichtyofauna of  Špičky u
Hranic na Moravě and Kelč localities. Part II: Kelč. Zpr. Vlas-
tivěd. Úst. v Olomouci 175, 1—9 (in Czech).

Knobloch E. 1969: Tertiäre Floren von Mähren. Moravské Museum

– Muzejní spolek, Brno, 1—201.

Kukalová-Peck J. 1991: Chapter 6: Fossil history and the evolution

of hexapod structures. In: Naumann I.D. (Ed.): The insects of
Australia, a textbook for students and research workers (2

nd

ed.).  Melbourne University Press, Melbourne, 141—179.

Martynov A. 1929: Fossil insects from Tertiary deposits in Ashutas

Saisan district. Trudy Geol. Muz. Akad. Nauk SSSR 5, 173—216.

Nel A., Martínez-Delcl

ès X., Paicheler J.-C. & Henrotay M. 1993:

Les ‘Anisozygoptera’ fossiles. Phylogénie et classification
(Odonata).  Martinia 3, 1—311.

Nel A., Martínez-Delcl

ès X., Escuillié F. & Brisac P. 1994: Les

Aeshnidae fossiles: Etat actuel des connaissances (Odonata,
Anisoptera).  Neu. Jb. Geol. Paläont.,  Abh. 194, 143—186.

Nel A., Petrulevičius J.F. & Jarzembowski E.A. 2005: New fossil

Odonata from the European Cenozoic (Insecta: Odonata:
Thaumatoneuridae, Aeshnidae, ?Idionychidae, Libellulidae).
Neu. Jb. Geol. Paläont.,  Abh. 235, 3, 343—380.

Prokop J. & Nel A. 2005: New scuttle flies from the Paleogene am-

ber in eastern Moravia of the Czech Republic (Diptera:
Phoridae).  Studia Dipterologica 12, 1, 13—22.

Prokop J., Nel A., Hájek J. & Bubík M. 2004: First record of a fos-

sil beetle (Coleoptera, Haliplidae) from the base of Paleocene
flysch sediments in the Magura Unit (Outer Western Car-
pathians, Moravia). Geol. Carpathica 55, 6, 469—473.

Přikryl T. 2004: New paleoichtyologic research at the Kelč locality

(Middle Oligocene, Czech Republic). In: Dostál O., Gregorová
R. & Ivanov M. (Eds.): Book of abstracts. 2

nd 

EAVP Meeting,

July 19—24. 2004, Brno, Czech Republic, 40.

Riek E.F. & Kukalová-Peck J. 1984: A new interpretation of drag-

onfly wing venation based upon Early Carboniferous fossils
from Argentina (Insecta: Odonatoidea) and basic characters
states in pterygote wings. Canad. J. Zool. 62, 1150—1166.

Říha P. 1979: Katalog der Tertiären und Quartären fossilen insekten

der Tsechoslowakei. Ent. Probl. (Bratislava) 15, 13—32.

Štys P. & Říha P. 1975: Two new genera and species of fossil Aly-

didae from the Tertiary of Central Europe (Heteroptera). Acta
Univ. Carolinae (Biol.) 1973, 185—197.

Sukatcheva I.D., Szalma Š., Vršanský P., Chalupová B., Golej M., La-

bajová E. & Smrečková M. 2006: Caddis-fly (Insecta: Tri-
choptera) from the Badenian volcano-sedimentary succession
(Western Carpathians, Slovakia). Geol. Carpathica 57, 531—534.

Wedmann S. 2000: Die Insekten der oberoligozänen Fossillagerstätte

Enspel (Westerwald, Deutschland). Systematik, Biostratinomie
und Paläoökologie. Mainzer Naturwiss. Archiv 23, 1—154.