Novák O (1877) Fauna der
Cyprisschiefer des Egerer Tertiärbeckens. Sitzungsberichte der
Koeniglichen Akademie der Wissenschaften,
Mathematisch-Naturwissenschaftliche Classe 76: 71-96.
ZooKeys 130: 299–305, doi: 10.3897/zookeys.130.1401
Ptychoptera deleta Novák, 1877 from the Early
Miocene of the Czech Republic: redescription of the first fossil
attributed to Ptychopteridae (Diptera)
Wieslaw Krzemiński1, Jakub Prokop2
1 Institute of Biology, Pedagogical University of Cracow, ul. Podbrzezie 3, 31-054 Kraków, Poland
2 Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44, Praha 2, Czech Republic
Abstract
The first fossil that was described in Ptychopteridae, Ptychoptera deleta Novák,
1877 from the classical Early Miocene locality Mokřina (Krottensee) in
western Bohemia is re-examined. The re-description of the holotype
including a new line drawing and remarks summarizing the scarce fossil
record of this group is provided.
KeywordsDiptera, Ptychopteridae,
Ptychoptera, revision, Tertiary, Neogene, Miocene, Ottnangian/Karpatian,
Cypris Formation, Cheb Basin, Czech Republic
Introduction
The family Ptychopteridae is a very small group with about 70 Recent species belonging to two subfamilies (Ptychopterinae and Bittacomorphinae), mostly distributed in the Holarctic, Ethiopian and Oriental Regions. In the Neotropics one species is currently known (Hancock et al. 2006). Ptychopteridae are not present in the Australian and Antarctic Regions. The oldest true representative, Ptychoptera mesozoica Kalugina, 1989 was described from the Lower Cretaceous (Neocomian) of Baissa in Buryatia (Siberia, Russia). Ptychoptera deleta
Novák, 1877, known from the Early Miocene of Mokřina (Krottensee) in
western Bohemia (Czech Republic), was the first described fossil
representative of the family. Handlirsch (1909) supposed that the species did not belong to the genus Ptychoptera Meigen, 1803 and created a new genus Ptychopterula mainly on the basis of a considerably narrow wing base as present in Etoptychoptera Handlirsch, 1909 and a free Sc. However, he emphasized also the common characters present in Ptychopterula and the Recent genera Ptychoptera and Bittacomorpha Westwood, 1835 connection of R and Rs, the position of crossvein r-m, etc. Alexander (1927) also supposed that the species described by Novák did not belong to Ptychoptera. Peus (1958), in his monograph on the Ptychopteridae, referred to this species as Liriope (“Ptychopterula”) deleta (Novák, 1877). These opinions were based on mistakenly Novak’s drawing only, without revision of the holotype (see Fig. 2B).
This paper presents a re-description of the holotype including a new
line-drawing and photograph. Five other compressed fossil ptychopterid
species were described till now: Ptychoptera miocenica (Cockerell, 1910) from the Oligocene of Florissant (Colorado, USA) originally placed in genus Bittacomorpha; Brodilka mitchelliLukashevich, Coram & Jarzembowski, 2001 and Zhiganka woolgariLukashevich, Coram & Jarzembowski, 2001 both from the Lower Cretaceous of Purbeck and Wealden groups (UK); Zhiganka comitans Lukashevich, 1995 from the Lower Cretaceous of Yakutia (Russia), and Probittacomorpha christenseni Freiwald & Willmann, 1992 from the lowermost Eocene of Mo-clay (Denmark) (Freiwald and Willmann 1992; Evenhuis 1994; Lukashevich 1995; Ansorge and Schröder 1999; Lukashevich et al. 2001). Two genera, ZhigankaandProbittacomorphaareattributed to the subfamily Bittacomorphinae and others to Ptychopterinae. A single species, Ptychoptera eocenica Podenas, 2007 was described from an Eocene Baltic amber inclusion (Podenas 2007).
Lukashevich (2008) proposed a new system of Ptychopteroidea and synonymized the family Eoptychopteridae known from the Upper Triassic to Lower Cretaceous with Ptychopteridae. Eoptychopteridae is a considerably variable group of flies with common occurrence throughout the Jurassic up to the Lower Cretaceous (Kalugina 1989; Krzemiński 1992; Lukashevich 1993; Lukashevich et al. 1998; Ren and Krzemiński 2002; Krzemiński and Krzemińska 2003).
Novák (1877) described a fossil entomofauna from Mokřina (Krottensee) located in Cheb county of western Bohemia (Czech Republic) (Fig. 1).
This classical Early Miocene locality in Cheb Basin yielded greenish
marls of lake sedimentation belonging to the Cypris Formation (Rojík 2004). Fossiliferous layers previously called “Cypris shales” are well known for abundant occurrence of ostracods e.g. Cypris angusta (Reuss, 1852) when Cheb and Sokolov basins were interconnected (Obrhelová and Obrhel 1987). Mokřina (Krottensee) locality belongs to biostratigraphic zone MN4 dated by mammals from nearby locality Dolnice (Fejfar 1974), and to ichthyozone IV defined by Obrhelová and Obrhel (1983). Fossil record from Mokřina is also well known by plants and bird remains (Bůžek et al. 1996; Mlíkovský 1996).
Novák’s collection from Mokřina housed in National Museum in Prague
contains 73 insect specimens classified in 10 families of five insect
orders (Prokop et al. 2003).
Figure 1.
AGeographical position of northwestern Bohemia within Europe B detailed map of the Cheb and Sokolov basins with indication of position of Mokřina (Krottensee) locality.
Material and methods
The holotype specimen was observed under stereomicroscope
Leica MZFLIII & MZ16 in dry state. The line drawing of the
venational pattern was drawn directly using a stereomicroscope and
finally readjusted to the photograph scale using image-editing software
(Adobe Photoshop CS). Photography was made simultaneously using a
digital camera Canon PowerShot S80 attached to the stereomicroscope. The
type material referred to as ‘NM’ is housed at the National Museum in
Prague, Czech Republic.
Systematic paleontology
Family Ptychopteridae Osten-Sacken, 1879
Genus
Ptychoptera
Meigen, 1803
Type species.
Tipula contaminata Linnaeus, 1758
Ptychoptera
deleta
Novák, 1877
http://species-id.net/wiki/Ptychoptera_deleta
Figs 2A–C
Ptychoptera deleta Novák, 1877: p. 88, Pl. II, Fig.1.
Ptychopterula deleta (Novák, 1877): Handlirsch (1909): p. 264, 269.
Liriope (“Ptychopterula”) deleta (Novák, 1878): Peus (1958):p. 12, fig. 17.
Diagnosis.
Rs very short; R4+5 five times longer than
Rs; R4 nearly as long as R4+5; wing coloration pattern with isolated
subapical spot and without spot on R2.
Redescription.
Wing with original coloration pattern of
dark clouds visible in medial and distal part; Sc rather long, ending
opposite proximal third part of R3; R1 long; cross vein r-r (R2) at its
about two lengths before tip of R1; Rs very short; R4+5 five times
longer than Rs; R4 nearly as long as R4+5; cross vein r-m just before
forks of Rs and of M; distal part R5 and most part of medial veins not
preserved, cross vein m-cu about its length behind origin of M3+4; A1
rather long, its distalmost section strongly curved to posterior wing
margin.
Figure 2 A–C.
Ptychoptera deleta Novák, 1877 A photograph of holotype specimen No. NM-P947 B original line drawing of wing venation from Novák (1877) C line drawing of wing venation (reconstruction) D Ptychoptera agnes Krzemiński and Zwick, 1993, drawing of wing venation (scale bar represents 1 mm).
Dimensions.
Length of wing fragment about 7 mm, probable total length about 7.1 mm; maximum width about 1.8 mm.
Holotype.
No. NM-P947 (imprint of nearly complete wing
with medio-apical part missing, posterior wing margin is partially
broken, venation well preserved with traces of original pattern of
coloration). Specimen is housed in collection of National Museum in
Prague, Czech Republic.
Age and layer.
Early Miocene (Ottnangian/Karpatian), Cypris Formation (grey claystone) sensu Rojík (2004).
Discussion.
We provide a re-description of the holotype
specimen with several inaccurate characters in wing venation corrected
in comparison to the original drawing of Novák, e.g., ending of R1 to
anterior wing margin, indication of coloration pattern, A1 basally
running parallel to Cu for long distance and ending in posterior wing
margin, several missing or present cross-veins. Moreover, we
supplement the description by the first photograph of this specimen
(unavailable in 1877).
Unfortunately the single diagnostic wing character (M1+2 fork) of the subfamily Ptychopterinae
is not preserved. However, the wing venation pattern with very well
visible crossvein r-m positioned just before M forking into M1+2 and
M3+4, and original color pattern of Ptychoptera deleta show considerable similarities to recent species of Ptychoptera. These are distinct apomorphic characters present only in this genus within the family Ptychopteridae. The wing venation of Ptychoptera deleta is somewhat similar to the recent Ptychoptera agnes Krzemiński & Zwick, 1993 described from Hungary (Figs 2C–D).
This fact is not surprising due to the Early Miocene age of our fossil
(about 18–16 Ma) and considerably high morphological stability of
insects (Nel and Prokop 2009; Hörnschemeyer et al. 2009).
Finally, our present re-examination of the holotype confirms the great
erudition of the Czech pioneer palaeontomologist Dr Ottomar Novák who
correctly attributed the fossil species to the otherwise modern genus Ptychoptera.
Acknowledgements
This paper is devoted to the 75th anniversary of Prof.
Alexandr Rasnitsyn, leader of Russian paleoentomological laboratory at
Paleontological Institute RAS, Moscow. Authors are grateful to Elena
Lukashevich, journal editor Dmitry Shcherbakov (both Palaeontological
Institute RAS, Moscow), and anonymous reviewer for their insightful
comments and suggestions to the early version of the manuscript. We are
grateful to Kamil Zagoršek (National Museum in Praha) for the loan of
the holotype specimen and Martin Fikáček (National Museum in Praha) for
his kind help with translation from German. Special thank goes to Iwona
Kania who kindly helped with scanning of line drawings to digital form.
Second author (JP) acknowledges the research support from Grant Agency
of the Czech Republic No. 205/08/0643 and Ministry of Education MSM
0021620828.
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