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New griffenfly, Bohemiatupus elegans from the Late Carboniferous of western Bohemia in the Czech Republic (Odonatoptera: Meganisoptera: Meganeuridae)

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A new griffenfly, Bohemiatupus elegans n. gen., n. sp. (Meganeuridae) is described from the Upper Carboniferous (Bolsovian) deposits of the Ovčín near Radnice in western Bohemia (Czech Republic). The new taxon based on fore- and hindwing venation is compared with the other meganeurid genera. It is the first record of a large griffenfly from the continental basins of the Bohemian Massif supplementing the other giant insects such as Bojophlebia prokopi Kukalová-Peck 1985 or Carbotriplura kukalovae Kluge 1996 from the same strata.
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Ann. soc. entomol. Fr. (n.s.), 2010, 46 (1–2) : 183-188
183
ARTICLE
New griff enfl y, Bohemiatupus elegans from the Late
Carboniferous of western Bohemia in the Czech Republic
(Odonatoptera: Meganisoptera: Meganeuridae)
Abstract. A new griffenfl y, Bohemiatupus elegans n. gen., n. sp. (Meganeuridae) is described
from the Upper Carboniferous ( Bolsovian) deposits of the Ovčín near Radnice in western Bohemia
(Czech Republic). The new taxon based on fore- and hindwing venation is compared with the other
meganeurid genera. It is the fi rst record of a large griffenfl y from the continental basins of the Bohemian
Massif supplementing the other giant insects such as Bojophlebia prokopi Kukalová-Peck 1985 or
Carbotriplura kukalovae Kluge 1996 from the same strata.
Résumé. Une nouvelle « libellule géante », Bohemiatupus elegans du Carbonifère supérieur de
la Bohème de l’ouest (République Tchèque) (Odonatoptera: Meganisoptera : Meganeuridae) .
Un nouveau genre et une nouvelle espèce de Meganeuridae (Meganisoptera), Bohemiatupus elegans
n. gen., n. sp. est décrit du Carbonifère supérieur d’Ovčín près de Radnice (ouest de la Bohême,
République Tchèque). Ce nouveau taxon est basé sur la nervation des ailes antérieures et postérieures.
Il est comparé aux autres Meganisoptera décrits. Il s’agit de la première « libellule géante » des bassin
continentaux du massif de Bohème, mais pas du premier insecte géant des mêmes aire et étage
(Bojophlebia prokopi Kukalová-Peck 1985, Carbotriplura kukalovae Kluge 1996).
Keywords: Taxonomy, Tupindae, Insecta, Bolsovian, Radnice Basin, Paleozoic.
J P (1),* & A N (2)
(1) Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 44, Praha 2, Czech Republic
(2) CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 rue Buff on, F-75005 Paris, France
* Corresponding author
E-mail: jprokop@natur.cuni.cz, anel@mnhn.fr
Accepté le 15 janvier 2010
Paleozoic odonatopteran order Meganisoptera
Martynov 1932 (so-called ‘Protodonata’) represents
the ‘stem group’ to true Odonata. However, the
members of Meganisoptera diff er from Odonata by
absence of nodus and pterostigma in wing venation,
as well as males lacking secondary genitalia (Bechly et
al. 2001). Above all, this group is well known for the
insects with largest wingspan that have ever lived such
as Meganeuropsis permiana Carpenter 1939 from Early
Permian of Kansas that reached wingspan more than
700 mm (Carpenter 1939, 1947).
e clade Odonatoptera is fi rst recorded about
320 million years ago in the Namurian, and its main
subgroups Odonatoclada Bechly, 2003 (= the clade
that comprises the Lapeyriidae Nel et al. 1999, the
Campylopteridae Tillyard 1928 and the Nodialata
Bechly 1996 viz. the Odonatoptera with a true nodus),
Meganisoptera, and Geroptera Brodsky 1994 suddenly
appeared in Westphalian (Brauckmann & Zessin 1989;
Riek & Kukalová-Peck 1984; Jarzembowski and Nel
2002). Meganisoptera are not very frequent in the fossil
record of Namurian and Westphalian deposits, viz.
the Westphalian Carpentertypus durhami (Carpenter
1960), Piesbergtupus hielscheri Zessin 2006, Arctotypus
diluculum (Whalley 1980), Palaeotherates pennsylvanicus
Handlirsch 1906, and Gallotupus oudardi Nel et al.
2008; and the Namurian Sinomeganeura huangheensis
Prokop et al. 2008, Shenzhousia qilianshanensis Zhang
et al. 2006, and Namurotypus sippeli Brauckmann &
Zessin 1989 (Zessin 2006; Zhang et al. 2006; Ren et
al. 2008; Nel et al. 2008, 2009).
e continental basins of the Bohemian Massif are
represented by a number of insect localities ranging
in age from the Duckmantian (Westphalian B) to
the Middle Stephanian. In the central and western
Bohemia the most famous localities are in the Radnice
Member of Bolsovian age (Wesphalian B-C), which
involves localities like Lubná, Příčina, Štílec, Vrapice
and Otvovice or in the Nýřany Member of the Asturian
age where Nýřany and Třemošná belong among the
most famous ones. Predominance of the Blattodea is
signifi cant in Nýřany and Lubná (Kušta 1883; Fritsch
1901; Handlirsch 1920; Kukalová 1955), while others
are represented by single records of palaeopterous
insects (Ephemeroptera, Paleodictyoptera), (Fritsch
1880; Kukalová 1958; Kukalová-Peck 1985; Novák
1880).
184
J. P & A. N
New insect material was recently discovered at
Ovčín near Radnice during extensive excavations by
palaeobotanists in tuff layers so called „whetstone
horizon“ from the roof of the Lower Radnice Seam
(Lower Bolsovian) with well known buried plants
preserved in situ (see Opluštil et al. 2007, 2009;
Libertín et al. 2009). Insect fauna is very rare with two
other undescribed specimens of palaeodictyopterid
larvae and orthopteroid insect (Prokop unpubl.).
Material and methods
e fossil specimen was observed under stereomicroscopes Zeiss
Cytoplast in dry state and under ethyl alcohol. Due to large
dimensions of the specimen the venation pattern line drawing
was redrawn from color photograph and revised by parallel
observation under stereomicroscope. Drawing was fi nally
readjusted to the photograph scale using of graphic software
(Adobe Photoshop). Photographs were made in dry state and
under ethyl alcohol by digital camera Nikon D80 with macro
lens Nikon AF-S VR Micro-Nikkor 105 mm in the highest
contrast as possible by single sided cross-light pre-exposure.
Type material was collected in 2006 during extensive excavations
by group of palaeobotanists so called “Czech mafi a” in Ovčin
near Radnice and housed in the collection of West Bohemian
Museum in Plzeň, Czech Republic.
We use the wing venation nomenclature of Nel et al. (1993),
and Bechly (1996). Systematic and divisions follows the concept
of higher phylogenetic classifi cation of Odonatoptera proposed
by Bechly (1996, 2007).
e venational symbols used here specifi ed as follows: symbols
in capitals denote the longitudinal veins (CA / CP – costal
anterior / posterior, ScA / ScP – subcostal anterior / posterior,
RA / RP – radial anterior / posterior, IR – intercalary radial
posterior, MA / MP – medial anterior / posterior, CuA / CuP
– cubital anterior / posterior, AA – anal anterior).
Figure 1
Bohemiatupus elegans n. gen., n. sp. (Meganeuridae), holotype M00485 (negative imprint) – line drawing of fore- and hindwing venation, isolated apical part
drawn as preserved.
New Late Carboniferous griff enfl y in western Bohemia
185
Results
Superorder Odonatoptera Martynov 1932
Order Meganisoptera Martynov 1932
Family Meganeuridae Handlirsch 1906
Subfamily Tupinae Handlirsch 1919
Genus Bohemiatupus n. gen.
Type species. Bohemiatupus elegans n. sp.
Etymology. Composite name after Bohemia (historical region
in central Europe, occupying the western two-thirds of the
traditional Czech Lands, currently the Czech Republic) and
Tupu s (genus name), masculine in gender.
Diagnosis. ScA short; ‘subnodus’ present between RA and RP
near base of RP2; parts of CuA and CuP between MP and AA
separated and oblique, with only one cells between them; AA1
with posterior branches; CuP very long and straight, with more
than three concave branches, but area covered by posterior
branches of CuP distinctly narrower than those of CuA and
AA1; hind wing distinctly broader and with anal and cubito-
anal areas distinctly broader than in forewing.
Bohemiatupus elegans n. sp.
(Figs. 1–4)
Holotype. Specimen M00485, well preserved basal and medial
part of fore- and hindwing, apical part of fore wing distorted
and medioapical part of hindwing not preserved, negative
imprint, collection of West Bohemian Museum in Plzeň, Czech
Republic.
Figures 2–4
Bohemiatupus elegans n. gen., n. sp. (Meganeuridae), holotype M00485 (negative imprint). 2, photograph of wing venation, ventral view; 3, detail photograph
of basal part of forewing; 4, detail photograph of forewing isolated apical part.
186
J. P & A. N
Derivation of name. Named after magnifi cent state of
preservation, elegans in Latin.
Type locality. Ovčín opencast mine near Radnice,
Radnice Basin, Czech Republic
Type strata. Upper Carboniferous, Westphalian B-C,
Bolsovian, Kladno Formation, Radnice Member, Low-
er Radnice Coal, Whetstone Horizon.
Description. Negative imprint of a fore and a
hindwing in life position, probably hyaline. Forewing
broadest part at about midwing; length of fragment
to fi rst distortion break about 115 mm, probable total
length about 260 mm; wing width 49 mm; estimate
ratio (wing length/width) about 5.3; precostal area of
CA and CP not preserved, but ScA is certainly short
as it is absent above arculus level; simple concave ScP
nearly parallel to anterior wing margin, and reaching it
in apical part of wing, at least distal of 190 mm from
wing base, but extreme apex of ScP not preserved; nine
visible ‘antenodal’ cross-veins in area between ScP and
C, more or less aligned with cross-veins in area between
ScP and RA; the most basal cross-vein perpendicular to
C, ScP, and RA, stronger than others ‘antenodals’, and
making a distinct brace corresponding to the primary
antenodal cross-vein Ax0 not preserved; ‘subnodus
between RA and RP near base of RP2; stem of radial
veins dividing 34 mm from wing base forming ‘arculus’,
RA strong, simple, parallel to ScP, and ending probably
in wing apex; R and MA fused in their basal parts; RP
bifurcating into RP1/2 and RP3/4 88 mm from wing
base; only basal part of RP3/4 preserved, simple; IR2
and other distal branches of RP partly preserved on
isolated apical part; MA nearly straight with its most
basal posterior branches partly preserved, fi rst opposite
bifurcation between RP1/2 and RP3/4; only fragments
of MP vein preserved and visible at level of arculus,
its medial part being hidden under rock, but MP
probably simple; at least CuA forming strongly oblique
cross-vein but remains of an oblique cross-vein CuP
also visible in area between MP and AA, CuP 15 mm
from wing base, CuA one cell distally; CuP separating
from CuA 36 mm from wing base and six cells distal of
CuP-crossing; distal part of CuA with numerous main
posterior branches; CuP very long and rather straight,
with four main simple posterior branches covering a
relatively small area, basal part before its fi rst furcation
long and straight, about 4 mm long, width of area of
CuP along posterior wing margin 51 mm; distal part
of AA forming a relatively large are, width of this are
along posterior wing margin 49 mm; basal part of AA
(+ Cu) forming a very weak curve; anal and cubito-
anal areas rather broad, 17 mm wide, with numerous
posterior branches.
e preserved parts of the hindwing are very
similar to the forewing in pattern of venation except
for the cubito-anal area, hindwing slightly broader
than forewing; length of wing fragment about 85 mm,
probable total length as about fore wing; wing width
50 mm; estimated ratio (wing length/width) 5.2; anal
and cubito-anal areas distinctly broader than in the
fore wing, 30 mm wide; preserved basal part of CuP
long and nearly straight but distal part not preserved;
distal part of AA forming a relatively large area with
8–9 main posterior branches.
Discussion
is Odonatoptera is clearly related to the
Meganisoptera that comprises the ‘families
Namurotypidae Bechly 1996, Meganeuridae
Handlirsch 1906, Kohlwaldiidae Guthörl 1962, and
Paralogidae Handlirsch 1906. Relationships with the
Namurotypidae are excluded because Bohemiatupus
n. gen. has a distally branched CuP, instead of being
simple.  e Paralogidae have a long and straight CuP as
in Bohemiatupus n. gen., but with numerous branches
covering a broader area than CuA and distal branch
of AA.  e Kohlwaldiidae are characterized by having
the distal parts of CuP and AA strongly reduced,
which is not the case of Bohemiatupus n. gen. (Nel et
al. 2009).  e Meganeuridae are characterized by the
presence of a typical ‘subnodal’ oblique vein between
RA and RP near the base of RP2, which is preserved
in our fossil.  e very large size of Bohemiatupus n.
gen. supports affi nities with this family even if several
Meganeuridae have distinctly smaller dimensions
(Nel et al. 2009). Bohemiatupus n. gen. cannot be
attributed to the Meganeurinae because its vein ScA is
short.  e Carpentertypinae Zessin 1983 [type genus
Carpentertypus Zessin 1983, based on one species
C. durhami (Carpenter 1961)] is based on a very
fragmentary fossil, with no information on any of the
basal halves of the wings. Nevertheless it diff ers from
Bohemiatupus n. gen. in its distinctly less numerous cells
and narrower area covered by MA.  e Piesbergtupinae
Zessin 2006 (type genus Piesbergtupus Zessin 2006) is
characterized by the narrow anal area of fore wing and
presence of one oblique cross-vein distal of the oblique
braces CuP and CuA and basal of the distal branch of
CuP re-emerging from CuA+CuP, plus the long CuP
and AA1 (Zessin 2006, 2008; Nel et al. 2009). e
pattern of wing venation of Bohemiatupus n. gen. ts
very well with the Tupinae.  e monophyly of this last
group is not very clearly established (Bechly 1996; Nel
et al. 2009), but we can now restrict the comparison of
Bohemiatupus n. gen. to the genera that are currently
attributed to it. After the key to the tupine genera of
Nel et al. (2009), Bohemiatupus n. gen. would fall
New Late Carboniferous griff enfl y in western Bohemia
187
near the Permian genus Tup us Sellards 1906 because
of the following characters: the two braces CuP and
CuA oblique, not fused into a long, oblique vein; AA1
with several posterior branches; and CuP with more
than three concave branches. Bohemiatupus n. gen.
greatly diff ers from Tupus in its hindwing cubito-anal
area distinctly broader than that of fore wing. Some
‘enigmatic’ genera are based on incomplete wings, i.e.
the Carboniferous genera Shenzhousia Zhang et al.
2006 and Boltonites Handlirsch 1919, and the Permian
Petrotypus Zalessky 1950. Petrotypus needs a complete
redescription, and it is based on a very poor fossil.
Nevertheless, its CuP seems to be simple and very
straight after Zalessky (1950), unlike Bohemiatupus n.
gen. Shenzhousia shares with Bohemiatupus n. gen. a
very long and straight CuP but with numerous posterior
branches covering a broad area, unlike Bohemiatupus
n. gen. (Zhang et al. 2006). After the fi gure of Bolton
(1922: fi g. 44), Boltonites has a distinctly curved CuP,
shorter than that of Bohemiatupus.
Conclusions
Bohemiatupus n. gen. is established for Bohemiatupus
elegans n. sp. (type species) that exhibits unique
combination of characters within Meganeuridae.  e
present discovery represent the fi rst record of large
griff enfl y from the continental basins of the Bohemian
Massif supplementing the other giant insects such
as Bojophlebia prokopi Kukalová-Peck 1985 or
Carbotriplura kukalovae Kluge 1996 from the same
strata.
Moreover, the material from Ovčín near Radnice
represents an extraordinary source of fossil plants in situ
and other animals like arachnid, fi sh and amphibian
trace fossils those enable study of developed peat
mire ecosystem into a shallow lake gradually fi lled by
redeposited volcanic ash from Bolsovian of western
Bohemia (Czech Republic).
Acknowledgements. e authors are grateful to Radek Labuťa
(National Museum in Praha) who discovered the present speci-
men and to group of the Czech paleobotanits so called “Czech
mafi a” for large excavations at Ovčín locality. We especially
thank to Josef Pšenička (West Bohemian Museum in Plzeň) for
the loan of the material.  e rst author (JP) acknowledges the
research support from the Grant Agency of Academy of the
Czech Republic No. KJB301110701 and Ministry of Schools
MSM 0021620828. We sincerely thank Dr Gunther Fleck for
his useful comments on the fi rst version of the paper.
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... Whereas lateral changes in the composition of plant fossil assemblages reflect spatial heterogeneity of vegetation cover, the vertical distribution of plant remains within a tuff includes separation of litter and ground cover from tree crown foliage, lianas and epiphytes (Burnham & Spicer 1986, Wing et al. 1993, Burnham 1994, Opluštil et al. 2009a. Evidence of interactions between animals and plants also may be preserved, not uncommonly (Prokop & Nel 2010, Scott & Taylor 1983, Rößler et al. 2012. ...
... The Whetstone Horizon bears coalified plant fossils and invertebrate remains (e.g. Opluštil et al. 2007, Prokop & Nel 2010. The character of preservation and distribution of fossil remains, however, significantly differs between the Bělka and Brousek. ...
... Most of the body fossils are arachnids (e.g. trigonotarbids; Fig. 24B) but also discovered in one of the previous excavations were the remains of a giant meganeurid griffenfly Bohemiatupus elegans (Prokop & Nel 2010; Fig. 24A), reaching about 55 cm of wing-span. In the S3 exacavation a few fragments resembling body segments of an Arthropleurid arthropod were found (Fig. 24C-E), although the exact affiliation of these fossils remains unknown. ...
Article
A Middle Pennsylvanian tuff bed (the Bělka bed) in the roof of the Lower Radnice Coal bears T0 peat-forming vegetation preserved in growth position. This vegetation, has been studied in detail at the 12 hectares large Ovčín coal deposit in the southern part of the Radnice Basin. Documentation of the fossil record in six excavations and that previously collected in the former opencast mine allowed for detailed reconstruction of the local peat-forming lepidodendrid–cordaitalean forest structured into well-developed storeys. It consists of about 33 species, which colonized the occasionally flooded planar peat swamp precursor of the Lower Radnice Coal. The canopy storey of this vegetation was dominated by Lepidodendron (Paralycopodites) simile, L. lycopodioides, Lepidophloios acerosus and Cordaites borassifolius. They formed a relatively dense canopy, locally interrupted with significant gaps allowing development of a rich groundcover that together with liana-like plants represents the most diverse part of the forest. A less diverse understory composed of calamites, medullosan pteridosperms and Psaronius tree ferns displays a patchy distribution pattern presumably related to density of the canopy. The minimal area that sufficiently represents the pattern of this forest phytocoenosis is estimated to be about 200 m2, although lower storeys are well represented even within much smaller areas of about 60 m2. Slight heterogeneity in the population density of dominant taxa (Cordaites vs. lepidodendrid lycopsids) was documented across the Ovčín coal deposit. The fossil record of the Bělka tuff bed also indicates that the coal-forest colonizing the peat swamp prior the generation of forest killed by volcanic ash fall, was destroyed, presumably due to long-lasting flooding and thus suggests that catastrophic events were probably a relatively common part of the evolution of peat-forming Pennsylvanian successions.
... The holotype of Carbotriplura kukalovae was found in the quarry 'Na Štilci' near Tlustice, Czech Republic, in tuffites of the Middle Pennsylvanian/Moscovian (Silesian, Westphalian C) (Kukalová-Peck, 1985). According to Prokop & Nel (2010), this deposit in the so-called Whetstone Horizon from the continental basins of the Bohemian massif represented a peat mire ecosystem with shallow lake that was gradually filled by re-deposited volcanic ashes from the Bolsovian of western Bohemia (Czech Republic). The Whetstone Horizon could be dated with the Ar/Ar method as 309 ± 3.7 Ma by Hess et al. (1985). ...
... Carbotriplura kukalovae was originally considered to exhibit an aquatic lifestyle, but our reinvestigation revealed that there are no obvious aquatic adaptations present in this fossil. The deposit holds both aquatic and terrestrial taxa (Prokop & Nel, 2010) as well as fossilized remnants of vegetation possibly over 20 m in height (see Opluštil et al., 2009). The presence of long legs with possibly multiple tarsal segmentation and large protruding eyes rather suggests that C. kukalovae had an arboreal, climbing lifestyle combined with diurnal activity. ...
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We revise the type material of the enigmatic fossil insect Carbotriplura kukalovae Kluge, 1996 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. C. kukalovae is regarded as a possible transitional fossil link between Zygentoma and Pterygota. Carbotriplurida is therefore elevated to ordinal rank and considered as putative fossil sister group of Pterygota. The paranotal theory of the origin of insect wings and the parachute theory of origin of insect flight are briefly discussed and further corroborated. Testajapyx thomasi from the Pennsylvanian of Mazon Creek is tentatively considered as Dermaptera rather than Diplura.
... The Carboniferous Odonatoptera include the well-known, large proto-odonates known as griffenflies (Meganisoptera: Meganeuridae) (Grimaldi and Engel 2005;Kohli et al. 2016), that had the largest wingspan (c. 71 cm) of all the insects that have ever lived (Prokop and Nel 2010;Nel et al. 2018). The Triassic was a pivotal period for Odonatoptera lineages (Jouault et al. 2022), with the diversification of the three groups 'Protozygoptera', Triadophlebiomorpha, and crown Odonata (Nel et al. 2012;Bechly 2016). ...
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The holotype is redescribed of the giant pterygote insect Bojophlebia prokopi Kukalová-Peck, 1985 from the Pennsylvanian of the Czech Republic. Multiple errors in the original description are documented and corrected. Bojophlebia prokopi has neither any visible traces of a costal brace nor an anal brace, but it does show triadic branchings of MA, MP, CuA, and even, as rare a plesiomorphy, of CuP. It is therefore rejected as a fossil stem mayfly and attributed as sister group of all other Hydropalaeoptera. The first cladistic analysis of fossil palaeopterous insects, including different palaeodictyopterid groups, is presented. A revised phylogeny of Hydropalaeoptera and the stem line of Ephemeroptera are suggested. Palaeodictyopterida is recognized as sister group of Neoptera; thus Palaeoptera s.l. is rejected as a paraphyletic taxon. Four new higher taxa - Paranotalia, Euhydropalaeoptera, Neopterygota and Litophlebioidea superfam. nov. - are introduced, as well as the new family Lithoneuridae.
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Gigantic as well as very large mayflies from the middle Upper Carboniferous (Westphalian) strata of Europe and North America are described: the adult and nymph of Bojophlebia prokopi n. gen., n. sp. (Bojophlebiidae n. fam.) and the nymphs of Lithoneura piecko n. sp. and Lithoneura clayesi n. sp. (Syntonopteridae). Evolution of ephemerid wing venation during 300 million years is summarized. Autapomorphic, apomorphic, and plesiomorphic character states of venation are categorized. Venational nomenclature of Recent Ephemerida is emended based on its evolutionary changes. Evidence that wing veins occurred primitively as a pair of fluted sectors is documented in Carboniferous mayflies in the costa, subcosta, radius, anal, and jugal. Ephemeroids and odonatoids are sister groups that share the veinal anal brace AA fused with CuP at an area important for flight. Ancestral Odonatoephemerida are the sister group of the extinct haustellate Paleoptera. The Carboniferous nymphs bear three pairs of almost homonomous th...
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A new dragonfly of family Meganeuridae Shenzhousia qilianshanensis gen. et sp. nov. discovered from Ningxia Hui Autonomous Region in North China is described in the present paper. It has an estimated wingspan of about 450-500 mm and may be the largest fossil insect in Late Carboniferous Namurian Stage discovered by far. The new species is referred to Meganeuridae because of the presence of the characteristic oblique vein between anterior branch of radius RA and posterior branch of radius RP near the base of RP2. It differs from other genera within the family in the following characteristics Precostal area short and not extending to the midwing posterior branch of subcostal vein short merging into costal vein near the level of originating point of IR2 RP forking earlier than anterior branch of media basally RP1+2 and RP3+4 parallel and close to each other for a long distance and then diverge gradually surpass midwing.
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The hitherto known members of the Odonata families „Eugeropteridae”, „Erasipteridae”, Paralogidae, Meganeuridae, and Triadotypidae are critically reviewed; their phylogenetic relationships are examined. Most important for these studies are three new finds from the Upper Namurian B from the famous insect bearing locality of Hagen-Vorhalle (Ruhr area, FRG). They represent a new genus and species of the Meganeuridae — Namurotypus sippeli n.g. n.sp. — and are the stratigraphically oldest known and most completely preserved specimens of this family as well. One of them for the first time shows the abdominal appendages of the male. For these — besides Erasipteroides valentini — most ancient unequivocally datable dragonflies at all the following features can be regarded as characteristic: (1) long antennae (more than twice as long as the head), (2) presumably rather small eyes, (3) small mandibles, (3) 4 tarsal segments (basitarsus + 3 tarsi), (5) unpaired median claw at the posttarsus, (6) only slightly oblique thoracic segments, (7) presence of 11 backwardly tapering abdominal segments, (8) presence of a terminal filum (as defined by Henning, 1968), (9) long paired penes at sternit 8, (10) large, leaf-like styli at the posterior end of sternit 8 (or perhaps to be interpreted as fused gonostyli + gonocoxites attached to sternit 9, respectively), (11) long cerci curved vertically like a reverse S, (12) obvious lack of accessory copulation apparatus at abdominal sternites 2 and 3, and (13) beginning of sclerotisation in the presubcostal area. Additionally, in strong contrast to the reconstruction of a meganeurid dragonfly (Meganeura monyi) by Handlirsch (1925), the fore wings do not overlap the hind wings. With a wing-span of 32 cm, Namurotypus sippeli is the largest known Namurian insect. These results on the morphology of Namurotypus sippeli allow a new and more appropriate reconstruction of the complete animal and furthermore lead to the following interpretation of the habit: (1) The animals probably were not very fast and agile during flight and mainly may have moved by gliding (with the abdominal appendages also operating as stabilizators); (2) secondary copulation is less probable; (3) it cannot be excluded that predatory feeding of the imagines was restricted. The critical review of the remaining taxa made it necessary to create three additional new genera: (1) Solutotherates n. g. with the type species S. analis (Carpenter, 1980) (incertae familiae) from the Westphalian D of Pennsylvania (USA), (2) Erasipteroides n. g. with the type species E. valentini (Brauckmann, 1985) (family „Erasipteridae”) from the Upper Namurian B of Hagen-Vorhalle (FRG), and (3) Whalleyala n. g. with the type species W. bolsoveri (Whalley, 1979) (family „Erasipteridae”) from the Westphalian A of Derbyshire (Great Britain). The results of the phylogenetic studies are summarized in a hypothetic dendrogram. Most probably the Paralogidae + Meganeuridae + Triadotypidae (= Meganisoptera sens. nov.) represent a monophyletic unit, whereas the „Eugeropteridae” and „Erasipteridae” are paraphyletic. Presumably the Meganisoptera as well as the recent Odonata have their origin in species of the „Erasipteridae”.
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The diagnosis of the Meganisoptera is discussed. Paralogus hispanicus sp. nov. is described from the Late Carboniferous-Early Permian of Spain. Those of the Paralogidae and Kohlwaldiidae and the positions of Oligotypus makowskii are discussed. The nomen nudum `Oligotypus britannicus' is transferred from Paralogidae into the Meganeuridae: Tupinae. Solutotherates is transferred in Kohlwaldiidae sit. nov. An emended diagnosis of Meganeuridae is proposed. The diagnoses of Meganeurinae Piesbergtupinae, and Tupinae are discussed and some important characters redefined. The description of Meganeura monyi is set forth. Meganeura (?) vischerae ZALESSKY, 1950, is considered as an Insecta incertae sedis star. nov. Meganeurula selysii and the genus Meganeurula are transferred into the Meganeurinae sit. nov. The genus Gilsonia MEUNIER, 1909, and the combination Gilsonia titana MEUNIER, 1909, are restored in Tupinae. The tupine genus Meganeurina HANDLIRSCH, 1919, is restored for Meganeurina confusa HANDLIRSCH, 1919. The diagnosis of the tupine type genus Tupus SELLARDS, 1906, is set forth, and a new species Tupus gallicus sp. nov. described from the Middle Permian of Lodeve. A new diagnosis is given for the genus Arctotypus MARTYNOV, 1932, with the following included species, which are either described or revised: four Russian species Arctotypus sinuatus MARTYNOV, 1932, Arctotypus sylvaensis MARTYNOV, 1940, Arctotypus fortis ZALESSKY, 1950, Arctotypus giganteus sp. nov., Arctotypus diluculum (WHALLEY 1980) comb. nov. from UK, and the four species from Lodeve Arctotypus gallicus sp. nov., Arctotypus merifonsensis sp. nov., Arctotypus intermedius sp. nov., and Arctotypus (?) undetermined species A. The new genus Permotupus gen. nov. is described for two new species Permotupus ollieorum sp. nov. and Permotupus minor sp. nov. from Lodeve. The new genus Curvitupus gen. nov. is described from Curvitupus ariegensis sp. nov. from Lodeve, Curvitupus verneti (LAURENTIAUX-VIEIRA & LAURENTIAUX 1963) comb. nov. from France, and two Russian species Curvitupus magnificus (ZALESSKY 1950) comb. nov. and Curvitupus elongatus (ZALESSKY 1950) comb. nov. The new genus and species Nannotupus pumilio gen. nov., sp. nov. is described from Lodeve. "Palaeotherates" HANDLIRSCH, 1906, and Ephemerites GEINITZ, 1865, are Meganisoptera incertae sedis stat. nov. Solutotherates BRAUCKMANN & ZESSIN, 1989, is transferred in the family Kohlwaldiidae sit. nov. The Lodeve griffenfly fauna is the most diverse throughout the entire Palaeozoic, with nine new species in five genera. It is also the most disparate in size with the smallest known species and very large taxa having wing sizes ranging from 50 mm long to more than 210 mm long, respectively. A key to the tupine genera is proposed.
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The first Carboniferous protozygopteran is formally described from the late Westphalian Coal Measures of southern England. Bechlya ericrobinsoni gen. et sp. nov. (Bechlyidae fam. nov.) is the oldest representative of a lineage which includes all living dragonflies and damselflies. This discovery shows that small, damselfly-like forms co-existed with the giant dragonflies of the Euramerican coal swamps.